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PROCEEDINGS

~ BIOLOGICAL SOCIETY ala |

_ WASHINGTON

VOLUME 111 NUMBER 3 18 SEPTEMBER 1998

ISSN 0006-324X

THE BIOLOGICAL SOCIETY OF WASHINGTON 1998-1999 Officers

President: Richard P. Vari Secretary: Carole C. Baldwin President-elect: Brian F. Kensley Treasurer: T. Chad Walter

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PROCEEDINGS

Editor: C. Brian Robbins

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

111(3):473—-503. 1998.

Horton H. Hobbs, Jr. (29 March 1914—22 March 1994). Biographical notes

Karen Reed and Raymond B. Manning

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A.

Abstract.—The 211 papers and abstracts published by the late Horton H. Hobbs, Jr., are cited and annotated by a list of new taxa published in each paper. An alphabetical list of taxa named by Hobbs is provided; the repository and the catalogue number are cited for each holotype. Hobbs’s reminiscences of his introduction to crayfishes and his early work are summarized from an

oral interview.

To many who knew him and his work, the name Horton H. Hobbs, Jr., (Figs. 1, 2) is synonymous with freshwater crayfishes of North America. Indeed, to many he was known affectionately as “‘crawdaddy.”’ The enormous advances in our understanding of the systematics, distribution, and evolution of crayfishes that resulted from his studies in a career spanning six decades are well known to all astacologists. Perhaps less known are his contributions on other groups, including freshwater and cave shrimps and crabs, and ostracod associates of crayfishes.

Hobbs’s studies resulted in the recogni- tion of many new taxa: | new family (Cam- baridae); 38 new genera and subgenera (1 genus of palaemonid shrimp, Neopalae- mon; 8 genera of entocytherid ostracods; and 29 genera and subgenera of North American crayfishes); and 286 species (168 crayfishes, 104 entocytherids, 8 caridean shrimps, and 6 freshwater crabs).

A remarkable number of Hobbs’s contri- butions are monographs or other book- length works on a wide variety of topics:

1942d, crayfishes of Florida, 179 pp.;

1964a, Hobbs & Villalobos, crayfishes of Cuba, 59 pp.;

1969a, Chace & Hobbs, freshwater and

terrestrial decapods of the West Indies, 258

PPp.;

1971b, entocytherid ostracods of Mexico and Cuba, 55 pp.;

1972a, subgenera of Procambarus, 22 PPp.;

1972g, crayfishes of North and Middle America, 173 pp.;

1974c, checklist of North and Middle American crayfishes, 161 pp.;

1977b, Hobbs & Peters, entocytherid os- tracods of North Carolina, 73 pp.;

1977c, Hobbs, Hobbs III, & Daniel, trog- lobitic decapods of the Americas, 183 pp.;

1981b, crayfishes of Georgia, 549 pp.; ~

1982e, Hobbs & Hart, revision of Atya, 143 pp.;

1986a, Andolshek & Hobbs, entocytherid ostracods of Georgia, 43 pp.;

1987b, revision of Astacoides, 50 pp.;

1989g¢, illustrated checklist of American crayfishes, 236 pp.

Hobbs’s productivity led a European stu- dent to remark to one of us (R.B.M.) “‘Oh, Hobbs. He writes faster than I can read.”’

Hobbs III recalls (in litt.) that during the late 1930s, Hobbs’s father held him by his ankles while Hobbs slithered into a vertical crack (River Bed Cave, Columbia County, Florida) to collect Procambarus (O.) palli- dus—he then pulled him back out (for other anecdotes by and about Hobbs, see Hobbs 1986d).

474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

7°

a

Fig. 1. Hobbs collecting after the 1967 meeting of the Association of Southeastern Biologists, Columbia, South Carolina (J. F Fitzpatrick, Jr. and Hobbs III).

J. F Fitzpatrick, Jr. (in litt.), provided two route on some field excursion. Because he other anecdotes. ““One day, in the ‘60s, I was in motion, he did not call or write think—Hobbs had stopped by to see Miss ahead to announce his arrival. When he Peggy on some business while he was en knocked on the door in the middle of the

VOLUME 111, NUMBER 3

day in an active residential neighborhood of a small Virginia city, he found Miss Peggy home alone. The entire conversation, not short, was conducted with him standing on the porch and she behind the cracked door with the chain engaged. To her, ‘It just wouldn’t be proper to have a man in the house with Miss Lucille away on an er- rand’, even such a true gentleman of long acquaintance as was Hobbs. He delighted in relating this story to those who knew her as an example of her propriety, but he never faulted her behavior; he only respected it in his amusement.”

“The other goes back to his graduate stu- dent days at Gainesville and reveals a side of his character that only those of us who knew him well knew. He and Archie Carr were classmates and shared an office with a few others. Despite the reticence of his later years, a youthful Archie apparently was animated and outspoken. He had a hab- it during repeated discussions of rearing back on the hind legs of his wooden chair, and when he made an emphatic point, he would slam the front legs on the floor. The mischievous crew connived to play a trick on Archie with Hobbs surely one of the conspirators if not the actual perpetrator. They got a couple of fireworks ‘torpedoes’ and clandestinely attached them to the front legs of Archie’s chair. When he came in, they led him into a topic to which they knew he would react strongly. True to form, he reared his chair and at an appropriate point slammed the front legs. Archie was attached to the ceiling by his nails as the others rolled on the floor in merriment. De- spite this treatment, the group remained friends for their whole lives.”

All of us who knew and worked with Horton have many fond remembrances of him. He was the quintessential southern gentleman, always rising when a woman entered his presence. It pained him not to be able to open doors for women, and his female car-poolers were trained to wait at a door if he fell behind them. Elizabeth Nel- son remembers that in his last years, in frail

475

condition and walking precariously with a cane, he rushed to open a door for her as she walked by with a small package. She knew that to refuse his offer of assistance would have pained him terribly.

Elizabeth also remembered his expres- sion, a mixture of embarrassment and delight, when he was presented with a pair of boxer shorts with two flies at his retire- ment party—the idea being that having studied entocytherid ostracods so long he might have developed hemipenes.

Brian Kensley joined Horton on one of his last field trips. “In the fall of 1987, I accompanied Horton on a field trip to east- ern Texas. He wanted to complete a survey of the crayfishes of this area, and also gath- er some data on Fallicambarus devastator Hobbs & Whiteman, 1987c, the burrowing crayfish that causes much destruction to fields and lawns. We drove in Horton’s car. He would set the speed control, and dis- course on the flora and geology of the area Wwe were driving through. Every now and then we would have to stop, so that he could push a net through a puddle next to the road, or to show me a tree with which I was unfamiliar. Between whiles, he re- counted incidents from his early days of field collecting and teaching. It was for me a natural history revelation. Arrived in Tex- as, we met with farmers and agricultural ex- tension officers to gather information on devastator, and to dig in the bone-dry, con- crete-hard fields dotted with chimneys. This was my primary role. Once I had got down to the water table, Horton would lie stretched out on the ground, one arm reach- ing 2—3 feet down into the hole for the cou- ple of crayfish at the bottom. We also did a lot of (much easier) digging in stream banks, and this is where I uncovered spec- imens of a beautiful spotted crayfish which he eventually named for me [Procambarus (Girardiella) kensleyi Hobbs, 1990a]. Hunt- ing season opened while we were in the Neches River area. On the third occasion while we were either walking through the woods, or working at a stream bank, and a

476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

VOLUME 111, NUMBER 3

rifle shot rang out nearby, Horton turned to me (I was distinctly edgy at this point) and said: “‘Let’s go home’, which we did, all the way back to northern Virginia, at the sedate speed limit while I absorbed yet more land- scape science.”

Horton thoroughly enjoyed his martini at cocktail hour (see Fig. 3, showing Horton on his boat, the martini barge, on Lake Bar- croft). While one of us (R.B.M.) was living in Tunisia, Horton graciously shared his home with me. On one of my trips back, I noticed that Horton was having a clear cocktail in a 12 oz. glass rather than in his normal martini glass. This appeared strange to me, but I didn’t comment on the large drink, assuming that Horton’s affinity for martinis had expanded exponentially. Someone told him of my concern, and he had many laughs from the occasion, as he had shifted from martinis to a gin and tonic, with lots of ice, for the summer. I suspect that I took Horton his last martini during his last stay in the hospital. It was in a glass jar, which he gleefully accepted and hid in a drawer so that he could have it at happy hour. Later that day, his daughter Nina vis- ited him, and Horton told her “‘look what I have.”’ She smelled the jar and asked where he got it, probably knowing full well that I was involved.

Aspects of Hobbs’s life and career have been summarized in other obituaries by Fitzpatrick (1995a, 1995b, 1996) and Hoff- man (1994). Hobbs’s southern manners were characteristic of him. One aspect of his life that was known to those who shared dinner as his guest was his interest (and ability) in baking and cooking. Alan David- son (1979:432) in North Atlantic Seafood published his recipe for hush puppies. Da-

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Fig. 2.

477

vidson referred to him as “The scholarly Mr Hobbs.”

Cooper & Cooper (1997:616) acknowl- edged Hobbs’s influence: ‘“‘As always, we are immensely grateful to the late Horton H. Hobbs, Jr., for the splendid lessons he taught (not all of them about decapods).”’

Only his colleagues in the Department of Invertebrate Zoology at the National Museum of Natural History are aware of one highlight of Hobbs’s career. In 1976 Thomas E. Bowman and Louis S. Kor- nicker, members of the department’s self- appointed SOL Awards Committee, awarded him the “‘Smithsonian Order of the Lobster.’’ The award celebrates an ac- complishment or achievement of outstand- ing insignificance, something that usually is quite difficult to associate with Hobbs. The award was based on the following se- ries of misadventures.

Hobbs and colleagues C. W. Hart, Jr. and Margaret Walton introduced four new names for the entocytherid ostracod, Don- naldsoncythere donnaldsonnensis (Klie, 1931) (see Hobbs & Peters 1977:43, 44 for a synonymy), already burdened by two synonyms, Entocythere humesi Hoff, 1943 and E. pennsylvanica Hart, 1960, as well as four unavailable names published as erroneous spellings: E. donnaldsoni Wolf, 1934-1938; E. donalsonensis Rioja, 1943; and E. donaldsonensis Tressler, 1947. Hobbs & Walton (1961a) named E. hiwasseensis from Georgia; Hart & Hobbs (1961b) named E. tuberosa from Tennes- see (with the erroneous spelling tubercu- lata in the same paper); Hobbs & Walton (1963b) named Donnaldsoncythere scalis from Virginia and D. ileata from Virginia.

United States crawfish workers, meeting of Association of Southeastern Biologists, Memphis, Ten-

nessee, 19 April 1969 (includes all significant workers with crayfishes and their associated biota, except for Perry C. Holt, who worked on branchiobdellids). From left to right, C. W. Hart, Jr., J. E Fitzpatrick, Jr., Glen Gentry, Rudolph Prins, Daniel J. Peters, James E Payne, Raymond W. Bouchard, Jean E. Pugh, Horton H. Hobbs III, Horton H. Hobbs, Jr, Marilyn Black, Martha Reiser Cooper, Joe B. Black, and John E. Cooper (J. E

Fitzpatrick, Jr., R. Prins, and Hobbs III).

THE BIOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 111, NUMBER 3

Biographical Notes

Here we reproduce part of an Oral His- tory Interview of Hobbs by Pamela M. Henson, Smithsonian Institution Archives, in July 1976. It and the rest of the inter- view, largely dealing with aspects of his ca- reer after he left the University of Florida to go to the University of Virginia in 1946, form Record Unit 9509 in the Smithsonian Institution Archives. The questions are by Henson, the answers by Hobbs.

Question: As we discussed, we can start with some biographical information about you and your education, and how you came to be interested in Crustacea.

Answer: My undergraduate, well, all of my work was done at the University of Florida. It was during the depression years, so when I received my Bachelor’s degree I was able to get an assistantship in the de- partment and took my Master’s. There was no place to go, so I stayed on and joined the faculty after I took my Master’s, teach- ing the introductory courses in biology. I stayed on for my doctorate; and it took me several years because of residence require- ments and working full time, so I got my degree in 1940. I spent one summer at the Mountain Lake Biological Station during that time, doing work with Dr. [Chauncy M.] Gilbert, a course in arthropods.

You asked about my interest in crayfish- es. This began when I was a freshman in college. We came to the laboratory exercise dealing with the crayfish; the first day it was external anatomy—had no difficulty whatsoever, everything was beautiful. The second day had to do with internal struc- tures, and the crayfish that I was given ap- parently had demised with the flood, be- cause it was complete soup inside. So I sup- pose I was an eager beaver, in part, and I knew that not far from the boarding house where I was living at the time there was a little creek. I went down after school that afternoon and I caught a few crayfishes and brought them back to the room. That eve- ning I made my dissection and everything

479

was as it should be, but I had some crayfish left over and I couldn’t see killing them. So I found a jar to put them in (I planned to take them back to the creek the next eve- ning) and put them on my desk where I was studying. In a few minutes I noticed a com- motion, and a male and female had gotten together. I watched them for a while, and I said, ‘““Well, maybe I’ll get eggs in a few days.’ And sure enough, within two weeks, eggs were there. About three weeks later the female was carrying the young, and the more I watched them the more interested I became. By the end of that semester, I went to the chairman of the department and told him that I’d become interested in crayfishes and would like to do some work with them. He said, “‘Well, I know nothing about them but I’ll be happy to help you in any way that I can.”’ So that’s the way it began, and since 1931, I’ve looked at not much else.

Question: That is interesting. And I sup- pose your room turned into a big aquarium?

Answer: No, not really. But strangely enough, I was never able after that time to rear a clutch of eggs to adulthood until after I came to the Smithsonian.

Question: Just chance?

Answer: I didn’t know enough about them at that time to do the wrong thing!

Question: That’s amazing. But there was no one at the university studying crayfish?

Answer: No, no, there was only one per- son in the United States at the time. The big crayfish men had died: [Walter] Faxon and [Arnold Edward] Ortmann; and Edwin P. Creaser at Michigan Museum [Museum of Zoology, University of Michigan] was the only person working.

That brings me to my first visit to the Smithsonian [Institution] which might be of some interest to you. In 1935, I went up to the Mountain Lake Biological Station for the summer. I had a letter of introduction from my major professor [J. Speed Rogers] at the University of Florida, and one from Dr. Ivey E Lewis, who was Dean of the University of Virginia, introducing me to Dr. Waldo [L.] Schmitt [then Head Curator

480

of Zoology at the U. S. National Museum]. So after my stay at the station, I came up to Washington hoping that I might be able to examine the crayfishes. Dr. Schmitt took one look at me ... I know you wouldn’t believe it now, at one time I looked a little young. At the ripe old age of twenty-one I suppose I looked like I was about fifteen. He told me that he would be glad to have me look at the crayfishes, but that I could not open any of the types, couldn’t open the bottles. Well I had made that trip at some sacrifice, and Dr. Creaser couldn’t identify my material.

Dr. Schmitt was very gracious but not of much help in doing so. Not until 1937, when I convinced Dr. Schmitt that I was a serious worker, did he allow me to come back and examine the type specimens. So I worked for five years on crayfishes not knowing the name of a single crayfish that I was working with!

Question: Oh my goodness! Yes, not having compared them with the ones here.

Answer: I needed to compare them with the types.

Question: Well, you must have come up with some original observations.

Answer: Oh, well, a number of my ani- mals were new. When I started working in Florida I think there’d been about, oh may- be, four species recorded from the state; and when I finished there were forty-two of them.

Question: [Laughter] That’s a difference!

Answer: Which labels me perhaps as a splitter.

Question: Yes, but still, there just hadn’t been that much work done. There was a paper written by Fenner [A.] Chace [Jr.] at one point about the lack of work in system- atics being done in that area during that pe- riod [reference to Chace 19517]. I guess it was not one of the more worked on areas?

Answer: No, it never has been. Crayfish- es have never attracted very many people. Those who start, most of them have fallen [by] the wayside after a little bit of work. I’ve worked on crayfishes I suppose longer

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

than any other human ever has, in terms of number of years spent. .

Question: What was the Smithsonian like when you first came here to work on those?

Answer: Oh, it was a delightful place. Of course, my coming from a small town and coming to Washington, in part, I suppose I was much impressed. Dr. Schmitt was most gracious, as were Mr. [Clarence R.] Shoe- maker, Mr. [James O.] Maloney, and Miss [Mary Jane] Rathbun.

Question: She was still here?

Answer: She was still here. I can’t re- member whether I met her or not, but at least I saw her. And Dr. [Leonhard] Stejne- ger was a marvel. All of them were most gracious and helpful, and as I said, I do not hold it against Dr. Schmitt at all because I suppose I did look like a high school kid who thought he was going to do something.

Question: Right, take apart all the type specimens.

Answer: With this particular group, the group that’s dominant in Florida, unless you look at the first pleopods of the males you can’t identify the crayfish at all, and through a bottle you couldn’t see them.

Question: Were the collections of cray- fish at that point fairly extensive here?

Answer: Well certainly nothing in com- parison with what we have at the present time. It was perhaps not even the most im- portant collection in the country. The Mu- seum of Comparative Zoology had many more types than we had at that time; and there were a few at the Philadelphia Acad- emy [of Natural Sciences], but a very im- portant collection existed here, one that formed a nucleus around which we’ ve been building on for a number of years. Now, of course, we have the largest collection of crayfishes in the world—perhaps larger than all the other collections of crayfishes in the world combined [the collection of crayfishes now includes about 25,000 lots and 1500 lots of types].

Question: Was anyone working on them then?

Answer: No, no one at the Smithsonian

VOLUME 111, NUMBER 3

had worked on crayfishes. I’m the first per- son.

Question: Yes, to come in and work on it.

Answer: Faxon worked at the Museum of Comparative Zoology, and Ortmann was at the Carnegie Museum, and Creaser was at the Michigan Museum. Perhaps you know that we have the Michigan collection now. After I came here my major professor be- came director of the Michigan Museum, and he said, ““Nobody’s working on cray- fishes out here, and we’re crowded. Wouldn’t the Smithsonian like to have it?” That was just before I came here; I was in Charlottesville at the time. He said, ““You’re nearby so you can use them from time to time.”’ So we inherited that collec- tion.

Question: How easy was it to start work- ing in a field like that where no one else was working?

Answer: As I look back on it, I had a delightful time all the way through. I knew that I could tell the difference between what I had, and just because I didn’t have a name for them at the time didn’t bother me too much. The literature had not been cluttered with my group of animals that I was work- ing on in Florida, so there was very little in the literature at all. So there was no great literature problem to face.

Question: Yes, going through the masses of descriptions.

Answer: I hate to say what has been done to it since. [Laughter.] As I frequently say, they will curse me and say all kinds of things, but they can’t ignore me any longer with the crayfish.

Question: Right, you have left your mark. Were there avenues, let’s say, for publishing your results?

Answer: Oh, they were very limited, of course. Again, I came along during the de- pression years, but Dr. Schmitt was very, very kind, and I can tell you one lovely little anecdote that happened to me in con- nection with publications. I used the Pro- ceedings of the Florida Academy of Sci-

481

ences—it used to be called the Journal. I used that for some of my work. Then the Charleston Museum at that time was pub- lishing a series of Leaflets and Zoologica out of New York, and The American Mid- land Naturalist [also were available]. So there were a number of places that were open. But I sent one of my early papers up to Dr. Schmitt, describing seven new cray- fishes from Florida—lI think it was seven new ones—and redescribing, in essence, a species that had already been described. In this I used the word “‘crawfish.”’ I'd always used crawfish as a name, and throughout most of the South they’re still referred to as crawfish. Dr. Schmitt carefully went through my manuscript completely and changed it to crayfish everywhere. So I de- cided that, well, if they’re going to force me to do this (I didn’t raise any questions whatsoever), Ill just accept it. So since that time I have used “crayfish,” and most of my students do. However, most of the other people, particularly those who’ ve worked in the South, still write crawfishes when they write it.

Question: 1 hadn’t even realized that there was the difference in words.

Answer: Oh yes. And one other thing oc- curred in that paper. I had caught crayfishes from a cave from which they had been re- ported in Florida. Faxon had identified the animal as Cambarus lucifugus [error for acherontis sensu Faxon, not Lonnberg; re- placed by lucifugus Hobbs (1940a)], a spe- cies that had been described by [Einar] Lonnberg in the latter part of the nineteenth century from a well down near Orlando. Well, believing that the written word was infallible, I assumed that the specimens from this cave were what Faxon said they were. This was in the same manuscript, and Dr. Schmitt passed it on up to Dr. Stejneger, who at that time was Head Curator. Dr. Stejneger took one look at it and he said, ‘“What right does he have to say that this is lucifugus?” [error for acherontis sensu Faxon.] He said, ““He should go down to that well and catch some crayfish from that

482

place to be sure.”” So Dr. Schmitt wrote back to me and gave me Dr. Stejneger’s re- marks. Well I was a little bit upset because I thought the possibility of my finding that well that had been dug back in the latter part of the last century was very slight, and Florida was not easy to get around in, it was certainly not like it is now—all highways. But a friend of mine, Lewis J. Marchand, who lived down near Orlando, happened to come by my office within a day or so after I'd heard from Dr. Schmitt. I asked him if he had ever seen any white crayfishes down in that area. He said, ““Oh sure, I know a spring where I’ve seen them a number of times. So I said, “How soon can you be ready to go?’’ And he said, “‘Well, tomor- row morning will be fine.’ So I said, “Well, we’ll start then.’’ Palm Springs is what it was. In the meantime, I had another friend, fellow student, at the university who was very good at water goggling—that’s what we called it in those days—it’s mod- ified scuba diving but you don’t have air and so on. Anyway, I asked him if he didn’t want to go. This was a cold November day, and, believe you me, Florida can get cold during the winter, the northern part. So we went down to Palm Springs, and when we got to this little spring I looked down—of course it hadn’t been used since summer— it was covered with algae, and lying on the algae were white crayfish everywhere. So this friend of mine who went along with us took his gear and jumped off into the spring and time after time he came up, so we got forty-four of those animals that day. I got back to the laboratory, and sure enough, it was not the same as the thing from the cave, and was precisely what Lonnberg had described. So I had to redescribe it and put a new name [Cambarus lucifugus]| on the material from the cave.

Question: Yes. You can’t ever count on

Answer: So Dr. Stejneger sitting in Wash- ington certainly saved me considerable em- barrassment, and taught me a good lesson not to accept the printed word.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Question: That is interesting, and you were having things published. Did you start collecting yourself at that point?

Answer: Oh, I started collecting back in 1931, when I was still a freshman.

Question: You did keep track of your dif- ferent types?

Answer: Oh yes. One of the biggest helps, I suppose, was Dr. [J. Speed] Rogers, my major professor and the chairman of the department at that time, with his meticu- lousness in keeping notes and insisting that everything be carefully curated. So I started out being trained as a curator, I suppose. Through the years I amassed a collection of some 80,000 specimens that I brought with me when I came to the Smithsonian. All of those were catalogued and we’re still using my old numbers. The collection’s so tre- mendous that we haven’t been able to in- corporate nearly all of them into the Smith- sonian catalogue, so we’re still using my old cards.

Question: Where did you keep it all?

Answer: In my basement at home.

Question: You did?

Answer: I had a tiny little office at the University of Virginia in an old building. The office was about the size of this little anteroom out here. In one of the rooms the floor fell in, the basement where I had put so many crayfish. I had to take everything out and have the floor reconstructed to sup- port the crayfish collection.

Question: Yes, I guess everything was pretty much alcoholic storage?

Answer: Everything was alcoholic stor- age, yes.

Question: Which is fairly heavy. The professors you were working under were they systematics people?

Answer: All of them were systematists, for the most part. There were four men in the department at the time, four of the full professors. All of them were graduates of the University of Michigan. One of them had taken his doctorate at, I believe, New York—I’m not sure whether it was Colum- bia [University] or not—but one of them

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had taken his doctorate elsewhere. But all of them had been trained in Michigan under [Alexander G.] Ruthven and [Robert W.] Hegner. Three of them were entomologists: Dr. Rogers worked on the Tipulidae, the crane flies; and Dr. [Theodore Huntington] Hubbell on the Orthoptera, grasshoppers, and Dr. [C. Francis] Byers on the Odonata, dragonflies. The fourth member was a mammalogist. He and I were good cronies because he was interested in bats—primar- ily, and bats live in caves and crayfish live in caves. So we had a delightful time team- ing up going on field trips.

I'll tell you one other story of Dr. [Harley Bakwel] Sherman, the mammalogist. The library at the University of Florida had got- ten a new photostat machine and had dis- carded the old one. Dr. Sherman and I res- cued it and rebuilt it. We had heard that there had recently been aerial photographs taken of the area around Gainesville, of the entire county, that were available in the county agent’s office. So we borrowed these, and made copies and placed them in our notebooks. Fortunately, those photo- graphs had been taken during the winter months when the deciduous trees had lost their leaves. This meant that where you would see a black spot on our maps this was a Cluster of live oak trees, and live oaks usually grow along some depression, ‘fre- quently indicating a sinkhole or maybe a cave. In Florida, that section of the state is quite flat and the roads are built on the sec- tion lines so that they run at mile intervals, almost straight, occasionally going around a sinkhole or something of the sort. But no place are you more than a half a mile from any spot—if you were on the road—from any place within the quadrangle. So we would ride down the section lines with our maps, and if we’d see a black spot, we'd get out of the car and look to see whether this was a cave in which there might be bats or crayfishes or something of the sort.

One afternoon we’d been out, it was get- ting quite late and there were two graduate students with us, one of them, Dr. [Jerome]

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Krivanek, who’s now at Vanderbilt [Uni- versity], and a young man, [William M.] McLane, who has recently died. Dr. Sher- man was driving, and I said, ““Dr. Sherman, here’s a little place right close to the road. Stop, it won’t take Billy and me a moment or two to have a look.”’ So we rushed over. It was nothing but a depression, but on our way back to the car there was a perfectly cylindrical chimney that went right down, dropped down, oh, between fifty and sev- enty feet. I peeked over the edge of it and saw that there was a little water in the bot- tom of it which excited me, so we rushed back to the car and asked them if we didn’t have time to make one quick drop into this hole to see what it was. McLane and I went down into the hole and the other two stayed up above. We got down to the bottom, and I saw white crayfish on the bottom. We looked around and there was a little open- ing into the side, about two and a half feet in diameter; and we crawled into this open- ing, and that led into a fissure that was about four feet wide and some sixty or sev- enty feet long, with no floor, but the entire thing with water under us. It was shallow at one end, then it dropped off rather quick- ly. Even to this day we don’t know how deep it really goes, it just continues on. I saw the white crayfish down there and I turned to this graduate student, and I said, “Billy, if you’ll catch one of those cray- fishes and it’s a new one, Ill name it for you.” Well, I knew what they were so I was perfectly safe. So Billy jumped into the wa- ter with a dip net and started scurrying around, but he was missing them. I said, ‘““Well, let’s go.’ The water had gotten so cloudy you couldn’t do anything. So we crawled back up to the top, and when we reached the top of the cave, he turned to me and said, ‘““Well, Doc, I didn’t get any of the big ones but I got this little one.”’ He handed me a vial with a little tiny crawfish [Troglocambarus maclanei; in his dedica- tion of this species to McLane, Hobbs (1942b:349, footnote) commented: “It is a pleasure to name this species for Mr.

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McLane, who has been a companion on many collecting trips, and who has added numerous valuable specimens to my collec- tion.’’] in it, the most amazing animal I had ever seen in my life. I couldn’t believe that it was anything except a mutant or some- thing had gone wrong somewhere.

We went on back into Gainesville, and I immediately got the binocular scope on it and saw that it was really something out of this world. So the next day I was anxious to get back and get some more of them. Most of the young men who usually went with me were tied up in classes, they couldn’t go. I found one man who’d go, and we had gone out to the car; we had our ropes and collecting equipment and so on. About that time I spotted the same man who had caught the crayfish down in Palm Springs, going across the campus. I called to him, and told him that we were going out to a cave, and wouldn’t he like to go, and he said no, that he had an organic exam the next day. ““Well, that’s too bad,”’ I said, “it’s a wonderful place to water goggle.” I knew that would get him, no question. “Oh?,”’ he said, “All right, Pll go.”’ I called him Jelly, so that gives you some idea as to his size. He wasn’t all that obese, he was pretty wide, he had some flesh on him and I was worried all this time about his getting through that side of the pit.

Question: That’s true, yes.

Answer: But, we got out to the cave, we went down it, . and he said, ““Now where’s that place to dive?’ I said, “It’s right through there.’’ He looked at the hole and he said, “I can make it.’’ So into the hole he went, all of us. It was such that we could prop our feet against one wall and lean our back against the other one, no place to stand whatsoever. He said, ‘““Now where’s that place to dive?”’ I said, ““You’re there.”’ Well, if looks could have killed any- one, well of course, I would have been dead. But he was a good sport so he said, ‘All right.”” So he put on his goggles, and he went down, and he came up time after time with this big white crayfish, Procam-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

barus pallidus, which I had recognized all along—but none of the little fellow. He said, ““Well, I’ve about had it.’’ I said, “‘Jel- ly, go down just one more time,”’ and this time when he came up he rotated just be- fore he surfaced and his light beam hit the submerged ceiling, and he saw one of them and picked it off and brought it up. After that, he went down and he got either three or four more that afternoon, all of them col- lected from the ceiling. So this little animal was highly adapted for living on submerged ceilings of caves; down below it would have been in competition, with a much less chance of survival certainly than it has above, a very small animal. What had hap- pened, of course, McLane, when he jumped into the water, had jarred the water so that the animal became dislodged and on the way down Billy happened to catch it.

Question: Going down without ever looking up. .

Answer: I finally caught one when Mar- chand, the man who had caught so many of them, jumped in one day and another one was dislodged and I saw it come loose and grabbed it with my dip net. That’s the only one I’ve ever caught.

Question: But it was, I guess, a com- pletely new type?

Answer: Oh, it was a completely new ge- nus [Troglocambarus]. It is the most fan- tastic crayfish that we know at the present time. Most crayfishes have teeth on their third maxilliped for chewing. In the first place the maxillipeds have tremendously large and long setae that interlock, and by carrying water through the gill chamber over this setal net, it filters its food out of the water. There is no other crayfish that utilizes this technique.

Question: Fascinating, too, that you did find it. Were there many students in system- atics at Florida at that time?

Answer: At that time, ecology was the thing, as it has become in recent years. I was trained as an ecologist; of course, the kind of ecology that we did then had little resemblance to the kind that exists at the

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present time. All of us were a combination systematist and ecologist. I was trained in ecology, and the only reason I got into sys- tematics was that I couldn’t identify my an- imals and there was no one to help me, and I had it to do.

Question: Yes, because that had not been done before. So you stayed there until 1946?

Answer: That’s right.

Question: And then you moved to the University of Virginia.

Answer: While I was there I came up here many weekends to work in the collec- tion.

Question: You were using these collec- tions?

Answer: Oh, yes, I was using them from the outset. At one time Dr. Schmitt was able to get a little funds to help me to come up and work on weekends. At that time, too, he did another very nice thing for me. Most of the people had to be out of the building by a certain hour, and I was permitted to stay on until midnight. Some way he man- aged to help me out to that extent, because it was so rare—I could only work on Sat- urday night. Sometimes I would come up on Friday, I could work Friday night and Saturday night, or on holidays when I came.

Question: I guess you got to know the collections here fairly well. Were they in fairly good condition—identified or cata- logued?

Answer: Oh, yes, all of the old material was, and much of the time that I was here, when I would come up on weekends, I was working up collections that had accumulat- ed and identifying them so that they could be catalogued by the time that I came back and they were very nice in sending me du- plicate cards for everything that I identified.

Question: Were there many collections coming in during those years, were there expeditions or collecting trips?

Answer: No, not a great many. There had been a considerable backlog because, as I said, no crayfish man had been here since [William Perry] Hay worked at the muse-

485

um. He also taught high school here in the Washington area, as I understand it. He’s one of the few older members—crayfish people—that I met, but he had retired and was quite an old man when I met him. He lived in Florida and made a special trip to come up to Gainesville to see me one time, so I was delighted. He subsequently gave me his library, so I have many of his old notes, things that will go to the archives eventually.

Co-authors of Horton H. Hobbs, Jr.

Hobbs published many papers with co- authors, acknowledging help in the field and collaborating with students and col- leagues. Here we list his co-authors and, where possible, their institution (usually as of the dates), to demonstrate the breadth of his association with others with similar in- terests. As pointed out by Hoffman (1994: 37) some of his papers are co-authored “sometimes for no other reason than to re- ward the collector of a new species.”

Andolshek, M. D. (1986a). Hobbs’s research assistant at the museum. National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. (see also Margaret A. Daniel).

Banner, Albert H. (1959d). Hawaii Institute of Marine Biology, University of Hawaii, Kaneohe.

Barr, Thomas C., Jr. (1960b, 1972b). University of Kentucky, Lexington.

Bedinger, M. S. (1964b, 1965a). U.S. Geological Sur- vey.

Bouchard, Raymond W. (1973a, 1976b, 1994). Acad- emy of Natural Sciences of Philadelphia.

Brown, Arthur V. (1987d). University of Arkansas, Fayetteville.

Burr, Brooks M. (1984b). Southern Illinois University, Carbondale.

Carlson, Paul H. (1983c, 1985a). Department of Health and Environmental Control, Columbia, South Car- olina.

Chace, Fenner A., Jr. (1959d, 1969a). National Mu- seum of Natural History, Smithsonian Institution, Washington, D.C.

Cooper, Martha R. (1972e, 1980c). North Carolina State Museum of Natural History, Raleigh.

Daniel, Margaret A. (1977c) (nee Margaret D. Andol- shek).

Fitzpatrick, Joseph EF, Jr. (1962d, 1970a, 1971d). Uni- versity of South Alabama, Mobile. Hobbs’s last stu- dent.

486

Franz, Richard (1983a, 1986b, 1991b, 1992). Florida State Museum, University of Florida, Gainesville. Freeman, Harry W. (1956d). College of Charleston,

South Carolina.

Grubbs, Andrew G. (1982b, 1986c). Southwest Texas State University.

Hall, Edward T., Jr. (1969d, 1972f, 1974d). Georgia Water Quality Control Board, Atlanta.

Hart, C. Willard, Jr. (1956e, 1959b, 1961b, 1966d, 1982e). National Museum of Natural History, Smithsonian Institution, Washington, D.C. One of Hobbs’s students.

Hobbs, Horton H., III (1962a, 1970c, 1973c, 1976e, 1977c, 1989e, 1990b, 1991d, 1995a, 1995b). Hobbs’s son, referred to herein as Hobbs III. Wit- tenberg University, Springfield, Ohio.

Holt, Perry C. (1967d, 1968b). Virginia Polytechnic Institute, Blacksburg. Hobbs’s first student. Fitzpat- rick recalls that Holt liked to refer to himself and Fitzpatrick as “alpha and omega.”

Hubricht, Leslie (1959d). Missouri Botanical Garden, St. Louis; Louisville, Kentucky.

Lee, David S. (1976c). North Carolina State Museum of Natural History, Raleigh.

Mackin, J. G. (1959d). Texas A&M University, Col- lege Station.

Manning, Raymond B. (1977d). National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C.

Marchand, Lewis J. (1943a). University of Florida, Gainesville.

Massmann, William H. (1952b). Virginia Fisheries Laboratory.

McClure, Auden C. (1983e). McLean, Virginia.

Means, D. Bruce (1972c). Tall Timbers Research Sta- tion, Tallahassee, Florida.

Page, Charles H. (1953b). Charlottesville, Virginia.

Parish, Claude E. (1949b). University of Alabama.

Penn, George Henry, Jr. (1958h). Tulane University, New Orleans, Louisiana.

Perkins, E O. (1967c). Virginia Institute of Marine Sci- ence, Gloucester Point.

Peters, Daniel J. (1977b, 1979a, 1982c, 1989b, 1991c, 1993). New Horizons Governor’s School for Science and Technology, Hampton, Virginia.

Pflieger, William L. (1988c). Fish and Wildlife Re- search Center, Missouri Department of Conserva- tion, Columbia.

Prins, Rudolph (1972d). Western Kentucky University, Bowling Green.

Robison, Henry W. (1982d, 1985b, 1988b, 1989f). Southern Arkansas University, Magnolia.

Rodriguez, Gilberto (1989c, 1989d). Instituto Vene- zolano de Investigaciones Cientfficas, Caracas.

Shoup, C. S. (1942c, 1947b). Vanderbilt University, Nashville, Tennessee.

Villalobos (Figueroa), (1958f,

Alejandro 1964a,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1974b, 1981a). Instituto de Biologia, Universidad Nacional Autonoma de México, México.

Walton sisters, Lucille (“Miss Lucille’’) and Margaret (“Miss Peggy”) (1957b, 1958d, 1959a, 1959c, 1960a, 1960c, 1961a, 1962b, 1963a, 1963b, 1966c, 1966e, 1967d, 1968b, 1968d, 1970b, 1971le, 1975b, 1976d, 1977f). Danville, Virginia and Mountain Lake Biological Station, Pembroke, Virginia. J. E Fitzpatrick, Jr. (in litt.) notes that ““They were—or at least Hobbs thought of them as—the quintessen- tial “Old Maid Schoolteachers.’ Both were older than he, Miss Lucille by a greater margin .... To the best of my knowledge they met at the Mountain Lake Biological Station where the sisters were sum- mer fixtures. They succumbed to his characteristic charm, and Miss Peggy, “who knew not a thing about crayfishes,”” took great satisfaction in contrib- uting to science by inking the pencil drawings. This seemed to be her contribution to the joint papers.”

Whiteman, Mike (1987c, 1991a). Texas Agricultural Extension Service, Lufkin.

Word, Benjamin H. (1958e). University of Virginia, Charlottesville. An undergraduate, now a M.D., who did a research project under Hobbs that was never published. At the time of his death Hobbs was pre- paring this report for publication; Hobbs III is pre- paring this final study (see also remarks at end of Hobbs’s bibliography).

Zinn, Donald J. (1948c). University of Rhode Island, Kingston.

Publications of Horton H. Hobbs, Jr.

Here we provide a complete bibliography of Hobbs’s publications, in chronological order, annotated with the names of new taxa in each publication. The citations are cross- referenced to the list of taxa named by Hobbs, given below. If the name of a taxon is given in the title, it is not repeated in the list of taxa named in that article.

1937. Some Florida crawfishes and their habitat dis- tribution. [Abstract]—Proceedings of the Florida Academy of Sciences for 1936 1:154.

1938a. Two new crawfishes from Florida. Cambarus hubbelli, Cambarus acherontis pallidus. [Ab- stract].—Proceedings of the Florida Academy of Sciences 2:90, 91. [Nomina nuda].

1938b. A new crawfish from Florida.—Journal of the Washington Academy of Sciences 28(2):61—65. Cambarus rogersi.

1940a. Seven new crayfishes of the genus Cambarus from Florida, with notes on other species.—Pro- ceedings of the United States National Museum 89: 387—423. C. hubbelli, C. kilbyi, C. lucifugus ala- chua, C. lucifugus lucifugus, C. pallidus, C. pictus, C. rathbunae.

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1940b. A new crayfish from South Carolina.—The Charleston Museum Leaflet 14:3—7. Cambarus lun- Zi.

1941a. A new crayfish from San Luis Potosi, México (Decapoda, Astacidae).—Zoologica, New York 26(1):1—4. Cambarus blandingii cuevachicae.

1941b. Three new Florida crayfishes of the subgenus Cambarus (Decapoda, Astacidae).—The American Midland Naturalist 26(1):110-121. C. byersi, C. cryptodytes, C. floridanus.

1942a. On the first pleopod of the male Cambari (De- capoda, Astacidae).—Proceedings of the Florida Academy of Sciences (for 1940) 5:55-61.

1942b. A generic revision of the crayfishes of the sub- family Cambarinae (Decapoda, Astacidae) with the description of a new genus and species.—The American Midland Naturalist 28(2):334—-357. Trog- locambarus, T. maclanei.

1942c. Hobbs, H. H., Jr., & C. S. Shoup. On the cray- fish collected from the Big South Fork of the Cum- berland River in Tennessee during the summer of 1938.—The American Midland Naturalist 28(3): 634-643.

1942d. The crayfishes of Florida.—University of Flor- ida Publication, Biological Science Series 3(2):179 pp.. pls. 1 (frontispiece), 2—24. Cambarellus schmit- ti, Procambarus apalachicolae, P. bivittatus, P. econfinae, P. escambiensis, P. geodytes, P. latipleu- rum, P. leonensis, P. okaloosae, P. pubischelae, P. Pycnogonopodus, P. pygmaeus, P. rogersi campes- tris, P. r. ochlocknensis, P. seminolae, P. shermani, P. youngi.

1943a. Hobbs, H. H., Jr., & L. J. Marchand. A contri- bution toward a knowledge of the crayfishes of the Reelfoot Lake area.—Journal of the Tennessee Academy of Science 18(1):6—35.

1943b. Two new crayfishes from the panhandle of Florida (Decapoda, Astacidae).—Proceedings of the Florida Academy of Sciences 6(1):49-—58. Note by Hobbs given in footnote (p. 56): ““Due to inadver- tent delay in publication of this volume diagnoses of these species [Procambarus leonensis, P. pyc- nogonopodus| appeared earlier in “The Crayfishes of Florida’ (Hobbs: Univ. Fla. Pub. Biol. Series 3(2): 114-115, 117) and thus actually constitute the orig- inal descriptions.”

1943c. Two new crayfishes of the genus Procambarus from Mexico (Decapoda, Astacidae).—Lloydia 6: 198-206. P. rodriguezi, P. toltecae.

1944. Notes on the subterranean waters of the Florida Peninsula with particular reference to their crusta- cean fauna.—The Biologist 26(1&2):6-8.

1945a. Notes on the first pleopod of the male Cam- barinae (Decapoda, Astacidae).—Quarterly Journal of the Florida Academy of Sciences 8(1):67—70.

1945b. The subspecies and intergrades of the Florida burrowing crayfish, Procambarus rogersi (Hobbs).—Journal of the Washington Academy of

487

Sciences 35(8):247—260. Note by Hobbs given in footnote (p. 260): “This paper was originally ac- cepted for publication in the Proceedings of the United States National Museum, and it was cited as ‘In press’ in my Crayfishes of Florida (Hobbs, 1942). Wartime restrictions, however, so delayed publication by the Museum that the manuscript was withdrawn and submitted to this JOURNAL in order that the full descriptions of the two new subspecies of Procambarus rogersi might appear more prompt- ly.”

1945c. Two new species of crayfishes of the genus Cambarellus from the Gulf coastal states, with a key to the species of the genus (Decapoda, Astacidae).— The American Midland Naturalist 34(2):466—474. C. diminutus, C. puer.

1947a. Two new crayfishes of the genus Procambarus from Georgia, with notes on Procambarus pubes- cens (Faxon) (Decapoda, Astacidae).—Quarterly Journal of the Florida Academy of Sciences 9(1):1— 18. P. enoplosternum, P. litosternum.

1947b. Hobbs, H. H., Jr, & C. S. Shoup. Two new crayfishes (Decapoda, Astacidae) from the Obey River drainage in Tennessee.—Journal of the Ten- nessee Academy of Science 22(2):138—-145. Cam- barus obeyensis, C. parvoculus.

1947c. A key to the crayfishes of the Pictus subgroup of the genus Procambarus, with the description of a new species from South Carolina.—The Florida Entomologist 30(3):25—31. P. lepidodactylus.

1947d. A preliminary report on the crayfishes of Vir- ginia. [Abstract].—Proceedings of the Virginia Academy of Science 1946—1947:72.

1948a. On the crayfishes of the Limosus section of the genus Orconectes (Decapoda: Astacidae).—Journal of the Washington Academy of Sciences 38(1):14— 21. O. shoupi.

1948b. Two new crayfishes of the genus Orconectes from Arkansas, with a key to the species of the Hy- las group (Decapoda: Astacidae).—The American Midland Naturalist 39(1):139-150. O. leptogono- podus, O. marchandi.

1948c. Hobbs, H. H., Jr, & D. J. Zinn. Crayfish in southern Nevada.—Science 107(2780):369.

1948d. The crayfish genus Cambarellus in the United States. [Abstract].—Proceedings of the Virginia Academy of Science 1948:88.

1948e. A new crayfish of the genus Orconectes from southern Tennessee (Decapoda, Astacidae).—Pro- ceedings of the Biological Society of Washington 61:85—91. O. wrighti.

1948f. A new crayfish of the genus Cambarus from Texas, with notes on the distribution of Cambarus fodiens (Cottle).—Proceedings of the United States National Museum 98:223-—231. C. hedgpethi.

1949a. The cave crayfishes of North America. [Ab- stract].—Journal of the Tennessee Academy of Sci- ence 24(3):170.

488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1949b. Hobbs, H. H., Jr, & C. E. Parish. Notes on the life history of a Virginia crayfish. [Abstract].—Pro- ceedings of the Virginia Academy of Science 1948— 1949:97.

1949c. Observations on the emergence of a stonefly of the genus Taeniopteryx in Virginia. [Abstract].— Proceedings of the Virginia Academy of Science 1948—1949:101.

1949d. A new crayfish of the genus Orconectes from the Nashville Basin in Tennessee, with notes on the range of Orconectes compressus (Faxon) (Decapo- da, Astacidae).—Proceedings of the Biological So- ciety of Washington 62:17—25. O. rhoadesi.

1950a. A new crayfish of the genus Cambarellus from Texas (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 63:89—94. C. ni- nae.

1950b. A new crayfish of the genus Procambarus from Oklahoma and Arkansas (Decapoda, Astacidae).— Journal of the Washington Academy of Sciences 40(6):194—198. P. tenuis.

1950c. Observations on the ecological distribution of three Virginia crayfishes. [Abstract]—The Virginia Journal of Science for 1949-1950, new series 1(4): 349.

1951a. A new crayfish of the genus Orconectes from southeastern Virginia (Decapoda, Astacidae).—The Virginia Journal of Science, new series 2(2):122— 128. O. virginiensis.

1951b. A new crayfish of the genus Procambarus from Louisiana, with a key to the species of the Spiculifer group.—Journal of the Washington Academy of Sci- ences 41(8):272—276. P. penni.

1952a. A new crayfish from Alabama, with notes on Procambarus lecontei (Hagen).—Proceedings of the United States National Museum 102:209-219. P. verrucosus.

1952b. Hobbs, H. H., Jr., & W. H. Massmann. The river shrimp, Macrobrachium ohione (Smith), in Virginia—The Virginia Journal of Science, new se- ries 3(3):206, 207.

1952c. A new crayfish of the genus Procambarus from Georgia with a key to the species of the Clarkii sub- group.—Quarterly Journal of the Florida Academy of Sciences 15(3):165—-174. P. howellae.

1952d. A new albinistic crayfish of the genus Cam- barus from southern Missouri with a key to the al- binistic species of the genus (Decapoda, Astaci- dae)—The American Midland Naturalist 48(3): 689-693. C. hubrichti.

1952e. A preliminary report on the crayfishes of the Atlantic Slope from New Brunswick to South Car- olina. [Abstract]——The Virginia Journal of Science, new series 3(4):295.

1953a. Two new crayfishes from the Highland Rim in Tennessee (Decapoda, Astacidae).—Journal of the Tennessee Academy of Science 28(1):20—27 [also published in Report of the Reelfoot Lake Biological

Station, vol. 17, 1953]. Cambarus brachydactylus, C. friaufi.

1953b. Hobbs, H. H., Jr, & C. H. Page. Additional records of the occurrence of the freshwater jellyfish, Craspedacusta sowerbii, in Virginia.—The Virginia Journal of Science, new series 4(3):137.

1953c. The epizootic associates of the crayfishes of the New River system with particular reference to the ostracods. [Abstract].—Journal of the Tennessee Academy of Science 28(3):180, 181.

1953d. A new crayfish of the genus Procambarus from Alabama and Florida (Decapoda, Astacidae).—Pro- ceedings of the Biological Society of Washington 66:173-178. P. suttkusi.

1953e. On the ranges of certain crayfishes of the Spi- culifer group of the genus Procambarus, with the description of a new species (Decapoda: Astaci- dae).—Journal of the Washington Academy of Sci- ences 43(12):412—417. P. raneyi.

1954a. Apparent competition between two groups of crayfishes in the southeastern states. [Abstract].— The Virginia Journal of Science, new series 4(4): 230.

1954b. A new crayfish from the upper coastal plain of Georgia (Decapoda, Astacidae).—Quarterly Journal of the Florida Academy of Sciences 17(2):110—118. Procambarus truculentus.

1954c. A redescription of Procambarus ruthveni (Pearse) from La Laja Creek at Cuatatotolapam, Ve- racruz, Mexico (Decapoda, Astacidae).—Occasional Papers of the Museum of Zoology, University of Michigan 559:1-5.

1954d. Studies on the geographic distribution of the crayfishes of the genus Procambarus. [Abstract].— Journal of the Tennessee Academy of Science 29(3): 181.

1954e. Notes on the evolution of the Longulus group of the crayfish genus Cambarus. [Abstract]—The Virginia Journal of Science, new series 5(4):261.

1955a. A new crayfish of the genus Cambarus from Mississippi.—Proceedings of the Biological Society of Washington 65:95—100. C. cristatus.

1955b. Ostracods of the genus Entocythere from the New River system in North Carolina, Virginia, and West Virginia.—Transactions of the American Mi- croscopical Society 74(4):325—333. E. daphnioides, E. runki.

1955c. A tendency towards cyclic dimorphism in fe- male crayfishes. [Abstract]—The Virginia Journal of Science, new series 6(4):248.

1955d. Two crayfish highways to Florida. [Ab- stract]_—Association of Southeastern Biologists Bulletin 2(1):7.

1956a. A new crayfish of the genus Cambarus from North Carolina and South Carolina (Decapoda, As- tacidae)—Journal of the Elisha Mitchell Scientific Society 72(1):61—67. C. reduncus.

1956b. A new crayfish of the Extraneus section of the

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genus Cambarus with a key to the species of the section (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 69:115-121. C. spicatus.

1956c. A new crayfish of the genus Procambarus from South Carolina (Decapoda: Astacidae).—Journal of the Washington Academy of Sciences 46(4):117— 121. P. echinatus.

1956d. Hobbs, H. H., Jr., & H. W. Freeman. The deca- pod crustaceans of the Wateree River system in North Carolina and South Carolina. [Abstract].— Association of Southeastern Biologists Bulletin 3(1):10.

1956e. Hart, C. W., & H. H. Hobbs, Jr. The crayfish of the Lower Flint-Chattahoochee River system. [Abstract]—The Virginia Journal of Science, new series 7(4):292.

1957a. Observaciones acerca de las especies del gé- nero Entocythere (Crustaceos, Ostracodos) de Cuba.—Anales del Instituto de Biologia, Universi- dad Nacional Aut6noma de México 27(2):431—436. E. hamata.

1957b. Hobbs, H. H., Jr, & M. Walton. Three new crayfishes from Alabama and Mississippi (Decapo- da: Astacidae).—Tulane. Studies in Zoology 5(3): 39-52. Procambarus hybus, P. jaculus, P. mancus.

1958a. The evolutionary history of the Pictus group of the crayfish genus Procambarus (Decapoda, Asta- cidae).—Quarterly Journal of the Florida Academy of Sciences 2(1):71—91.

1958b. Two new crayfishes of the genus Procambarus from South Carolina.—Journal of the Washington Academy of Sciences 48(5):160—168. P. ancylus, P. hirsutus.

1958c. Two new crayfishes of the genus Procambarus from South Carolina and Georgia.—Notulae Natu- rae, Academy of Natural Sciences of Philadelphia 307:1—10, pls. 1, 2. P. chacei, P. epicyrtus.

1958d. Hobbs, H. H., Jr, & M. Walton. Procambarus pearsei plumimanus, a new crayfish from North Car- olina (Decapoda, Astacidae).—Journal of The Eli- sha Mitchell Scientific Society 74(1):7—12.

1958e. Word, B. H., & H. H. Hobbs, Jr. Observations on the testis of the crayfish Cambarus montanus ac- uminatus Faxon.—Transactions of the American Microscopical Society 77(4):435—450.

1958f. Hobbs, H. H., Jr, & A. Villalobos. The exo- skeleton of a freshwater crab as a microhabitat for several invertebrates. [Abstract].—The Virginia Journal of Science, new series 9(4):395, 396.

1958g. General Zoology. Tracey I. Storer & Robert L. Usinger. [Book review].—American Institute of Bi- ological Sciences Bulletin 8:43.

1958h. Penn, G. H., & H. H. Hobbs, Jr. A contribution toward a knowledge of the crawfishes of Texas (De- capoda, Astacidae)—The Texas Journal of Science 10(4):452-—483.

1959a. Hobbs, H. H., Jr., & M. Walton. A new crayfish

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of the genus Procambarus from Alabama (Decap- oda, Astacidae).—Proceedings of the Biological So- ciety of Washington 72:39—44. P. lewisi.

1959b. Hobbs, H. H., Jr., & C. W. Hart, Jr. The fresh- water decapod crustaceans of the Apalachicola drainage system in Florida, Southern Alabama, and Georgia.—Bulletin of the Florida State Museum, Bi- ological Series 4(5):145—191. Procambarus rogersi expletus.

1959c. Walton, M., & H. H. Hobbs, Jr. Two new eye- less ostracods of the genus Entocythere from Flori- da.—The Quarterly Journal of the Florida Academy of Sciences 22(2):114—120. E. ambophora, E. luci- fuga.

1959d. Hobbs, H. H., Jr., E A. Chace, Jr., J. G. Mackin, L. Hubricht, & A. H. Banner. Malacostraca. Pp. 889-901 in H. B. Ward & G. C. Whipple (W. T. Edmondson, editor), Freshwater Biology, 2nd edi- tion, Wiley, New York, 1248 pp. [crayfishes, pp. 882-898].

1960a. Hobbs, H. H., Jr, & M. Walton. Three new ostracods of the genus Entocythere from the Hiwas- see drainage system in Georgia and Tennessee.— Journal of the Tennessee Academy of Science 35(1): 17-23. E. cyma, E. mecoscapha, E. simondsi.

1960b. Hobbs, H. H., Jr, & T. C. Barr, Jr. The genus Cambarus. The origins and affinities of the troglob- itic crayfishes of North America (Decapoda, Asta- cidae), I—The American Midland Naturalist 64(1): 12—33. C. jonesi.

1960c. Hobbs, H. H., Jr., & M. Walton. A new crayfish of the genus Procambarus from southern Alabama (Decapoda, Astacidae).—Proceedings of the Biolog- ical Society of Washington 73:123—129. P. lophotus.

1961a. Hobbs, H. H., Jr., & M. Walton. Additional new ostracods from the Hiwassee drainage system in Georgia, North Carolina, and Tennessee.—Transac- tions of The American Microscopical Society 80(4): 379-384. Entocythere falcata, E. hiwasseensis.

1961b. Hart, C. W., Jr, & H. H. Hobbs, Jr. Eight new troglobitic ostracods of the genus Entocythere (Crustacea, Ostracoda) from the eastern United States.—Proceedings of The Academy of Natural Sciences of Philadelphia 113(8):173-185. E. arcu- ata, E. barri, E. pholetera, E. prionata, E. steevesi, E. tuberosa, E. ungulata, E. xania.

1962a. Hobbs, H. H., Jr, & H. H. Hobbs Ill. A new crayfish of the genus Cambarus from Georgia (De- capoda, Astacidae).—Proceedings of the Biological Society of Washington 75:41—45. C. conasaugaen- SiS.

1962b. Hobbs, H. H., Jr, & M. Walton. New ostracods of the genus Entocythere from the Mountain Lake Region, Virginia (Ostracoda, Entocytheridae).—The Virginia Journal of Science, new series 13(2):42—48. E. asceta, E. chalaza, E. phyma.

1962c. Notes on the affinities of the members of the Blandingii section of the crayfish genus Procam-

490

barus (Decapoda, Astacidae).—Tulane Studies in Zoology 9(5):273—293.

1962d. Hobbs, H. H., Jr., & J. FE Fitzpatrick, Jr. A new crayfish of the Propinquus group of the genus Or- conectes from the Ohio system drainage in West Virginia (Decapoda: Astacidae).—Proceedings of the Biological Society of Washington 75:207—214. O. propinquus erismophorous.

1962e. La presencia de Procambarus clarkii (Girard) en los estados de Chihuahua y Sonora, México (De- capoda, Astacidae).—Anales del Instituto de Biol- ogia, Universidad Nacional Aut6noma de México 33(1 &2):273—276.

1963a. Hobbs, H. H., Jr., & M. Walton. Three new ostracods (Ostracoda, Entocytheridae) from the Duck River drainage in Tennessee.—The American Midland Naturalist 69(2):456—461. Ankylocythere hyba, Dactylocythere xystroides, Uncinocythere zancla.

1963b. Hobbs, H. H., Jr., & M. Walton. Four new spe- cies of the genus Donnaldsoncythere (Ostracoda, Entocytheridae) from Virginia with a key to the spe- cies of the genus.—Transactions of the American Microscopical Society 82(4):363—-370. D. ardis, D. ileata, D. scalis, D. truncata.

1963c. Florida crayfishes.—The Quipu 1(2):8.

1964a. Hobbs, H. H., Jr, & A. Villalobos. Los cam- barinos de Cuba.—Anales del Instituto de Biologia, Universidad Nacional Autonoma de México 84(1 &2):307—366. Procambarus niveus.

1964b. Hobbs, H. H., Jr, & M. S. Bedinger. A new troglobitic crayfish of the genus Cambarus (Decap- oda, Astacidae) from Arkansas with a note on the range of Cambarus cryptodytes Hobbs.—Proceed- ings of the Biological Society of Washington 77:9— 15. C. zophonastes.

1964c. A new cave-dwelling crayfish from the Green- brier drainage system, West Virginia (Decapoda, As- tacidae).—Proceedings of the Biological Society of Washington 77:189—194. Cambarus nerterius.

1965a. Bedinger, M. S., & H. H. Hobbs, Jr. Observa- tions of a new troglobitic crayfish (with notes on the distribution of troglobitic crayfishes in the Ozark Region).—National Speleological Society, Bulletin 27(3):93-96.

1965b. A new crayfish of the genus Cambarus from Tennessee with an emended definition of the genus (Decapoda, Astacidae).—Proceedings of the Biolog- ical Society of Washington 78:265—-273. C. pristin- us.

1966a. Astacus oreganus Randall, 1840 (Crustacea, Decapoda): proposed suppression under the Plenary Powers. Z.N.(S.) 1727.—Bulletin of Zoological No- menclature 22(5&6):351—354.

1966b. A new crayfish from Alabama with observa- tions on the Cristatus section of the genus Cambarus (Decapoda, Astacidae).—Proceedings of the Biolog-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ical Society of Washington 79:109-116. C. promi- nens.

1966c. Hobbs, H. H., Jr, & M. Walton. Orconectes juvenilis (Hagen) in Mountain Lake, Virginia: an unplanned experiment in interspecific competition (Decapoda, Astacidae)—The Virginia Journal of Science, new series 17(3):136—140.

1966d. Hobbs, H. H., Jr, & C. W. Hart, Jr. On the entocytherid ostracod genera Ascetocythere, Plec- tocythere, Phymocythere (gen. nov.), and Cymocy- there, with description of new species.—Proceed- ings of The Academy of Natural Sciences of Phil- adelphia 118(2):35-61. A. coryphodes, A. cosmeta, A. didactylata, A. hoffmani, A. hyperoche, A. my- xoides, A. ozalea, A. sclera, C. gonia, Plectocythere Johnsonae.

1966e. Hobbs, H. H., Jr., & M. Walton. A new genus and six new species of entocytherid ostracods (Os- tracoda, Entocytheridae).—Proceedings of the Unit- ed States National Museum 119:1—12. Dactylocy- there brachystrix, D. pachysphyrata, D. cayugaen- sis, Entocythere kanawhaensis, Thermastrocythere, T. harti, Unicinocythere stubbsi.

1966f. An illustrated key to the species of the genus Ankylocythere with a description of a new species from Louisiana (Ostracoda, Entocytheridae).—The Proceedings of the Louisiana Academy of Sciences 29:67-75. A. harmani.

1967a. A new genus and three new species of ostra- cods with a key to genus Dactylocythere (Ostracoda: Entocytheridae).—Proceedings of the United States National Museum 122:1—10. D. jeanae, D. phoxa, Ornithocythere, O. waltonae.

1967b. The current status of the crayfishes listed by Girard (1852) in his “‘A revision of the North Amer- ican Astaci ...’’ (Decapoda, Astacidae).—Crusta- ceana 12:124—132.

1967c. Hobbs, H. H., Jr, & E O. Perkins. A new bur- rowing crayfish from North Carolina (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 80:141—146. Cambarus catagius.

1967d. Hobbs, H. H. Jr, P. C. Holt, & M. Walton. The crayfishes and their epizootic ostracod and bran- chiobdellid associates of the Mountain Lake, Vir- ginia, Region.—Proceedings of the United States National Museum 123:1—84.

1967e. A new crayfish from Alabama caves with notes on the origin of the genera Orconectes and Cam- barus (Decapoda: Astacidae).—Proceedings of the United States National Museum 123:1—17. Procam- barus pecki.

1968a. Two new crayfishes of the genus Cambarus from Georgia, Kentucky, and Tennessee (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 81:261—274. C. halli, C. sphenoides.

1968b. Hobbs, H. H., Jr., BR C. Holt, & M. Walton. The crayfishes and their epizootic ostracod and bran- chiobdellid associates of the Mountain Lake, Vir-

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ginia, region. [Abstract]—The Virginia Journal of Science, new series 19(3):178.

1968c. Crustacea: Malacostraca. Pp. K1—-K36 in E K. Parrish, Keys to water quality indicative organisms (southeastern United States). Federal Water Pollu- tion Control Administration, U.S. Department of In- terior.

1968d. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods from the southern United States.— Proceedings of the Academy of Natural Sciences of Philadelphia 120(6):237—252. Ascetocythere batchi, Dactylocythere cooperorum, D. prinsi, Entocythere reddelli, Harpagocythere tertius, Litocythere, L. lu- cileae, Uncinocythere warreni.

1969a. E A. Chace, Jr., & H. H. Hobbs, Jr. The fresh- water and terrestrial decapod crustaceans of the West Indies with special reference to Dominica.— United States National Museum Bulletin 292:258 Pp.

1969b. Procambarus villalobosi, un nuevo cambarino de San Luis Potosi, México (Decapoda, Astaci- dae).—Anales del Instituto de Biologia, Universidad Nacional Aut6énoma de México, Serie Ciencias del Mar y Limnologia, 38(1)(for 1967):41—46.

1969c. On the distribution and phylogeny of the cray- fish genus Cambarus. Pp. 93-178 in P. C. Holt, R. L. Hoffman, & C. W. Hart, Jr, eds. The distribu- tional history of the biota of the southern Appala- chians, Part I: Invertebrates. Virginia Polytechnic In- stitute, Research Division Monograph 1:295 pp. Av- iticambarus, Barbicambarus, Depressicambarus, Erebicambarus, Fallicambarus, Jugicambarus, Lacunicambarus, Veticambarus.

1969d. Hobbs, H. H., Jr, & E. T. Hall, Jr. New cray- fishes from Georgia (Decapoda Astacidae).—Pro- ceedings of the Biological Society of Washington 82:281—294. Cambarus howardi, C. unestami.

1969e. Two new species of the crayfish genus Procam- barus (Decapoda, Astacidae) with keys to the mem- bers of the Spiculifer group.—Proceedings of the Biological Society of Washington 82:329-348. P. elegans, P. gibbus.

1970a. Hobbs, H. H., Jr., & J. E Fitzpatrick, Jr. A new crayfish of the genus Fallicambarus from Tennessee (Decapoda, Astacidae).—Proceedings of the Biolog- ical Society of Washington 82:829—836. F. hortoni.

1970b. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods from Tennessee and Virginia.—Pro- ceedings of the Biological Society of Washington 82:851—863. Ascetocythere holti, Dactylocythere en- oploholea, D. myura, D. spinata.

1970c. Hobbs, H. H., Jr, & H. H. Hobbs III. New entocytherid ostracods with a key to the genera of the subfamily Entocytherinae.—Smithsonian Con- tributions to Zoology 47:19 pp. Ascetocythere lita, Dactylocythere coloholca, D. macroholca, D. pug-

Hiaticambarus, Puncticambarus,

491

hae, Entocythere tyttha, Geocythere nessoides, Lor- docythere, L. petersi.

1970d. A new crayfish from the Nashville Basin, Ten- nessee.—Proceedings of the Biological Society of Washington 83:161—169. Cambarus gentryi.

1970e. New crayfishes of the genus Cambarus from Tennessee and Georgia (Decapoda, Astacidae).— Proceedings of the Biological Society of Washing- ton 83:241—259. C. bouchardi, C. cymatilis.

1971a. A new crayfish of the genus Procambarus from Mississippi (Decapoda: Astacidae).—Proceedings of the Biological Society of Washington 83:459—468. P. fitzpatricki.

1971b. The entocytherid ostracods of Mexico and Cuba.—Smithsonian Contributions to Zoology 81: 55 pp. Ankylocythere maya, A. toltecae, A. villalo- bosi, Uncinocythere zaruri.

1971c. New crayfishes of the genus Procambarus from Alabama and Texas (Decapoda, Astacidae).—Pro- ceedings of the Biological Society of Washington 84:81—94. P. capillatus, P. texanus.

1971d. Fitzpatrick, J. E, Jr., & H. H. Hobbs, Jr. A new crawfish of the Spiculifer group of the genus Pro- cambarus (Decapoda, Astacidae) from central Mis- Sissipp1.—Proceedings of the Biological Society of Washington 84:95—-102. P. lylei.

1971le. Walton, M., & H. H. Hobbs, Jr. The distribution of certain entocytherid ostracods on their crayfish hosts.—Proceedings of the Academy of Natural Sci- ences of Philadelphia 123(4):87—103.

1971f. A new troglobitic crayfish from Florida.— Quarterly Journal of the Florida Academy of Sci- ences 34(2):114—-124. Procambarus milleri.

1972a. The subgenera of the crayfish genus Procam- barus (Decapoda: Astacidae).—Smithsonian Contri- butions to Zoology 117:22 pp. Acucauda, Austro- cambarus, Capillicambarus, Hagenides, Leconti- cambarus, Lonnbergius, Mexicambarus, Pennides, Remoticambarus, Scapulicambarus, Tenuicamba- rus, Villalobosus.

1972b. Hobbs, H. H., Jr., & T. C. Barr, Jr. Genus Or- conectes. Origins and affinities of the troglobitic crayfishes of North America (Decapoda, Astacidae), II.—Smithsonian Contributions to Zoology 105:84 pp. O. incomptus.

1972c. Hobbs, H. H., Jr, & D. B. Means. Two new troglobitic crayfishes (Decapoda, Astacidae) from Florida.—Proceedings of the Biological Society of Washington 84:393—409. Procambarus horsti, P. orcinus.

1972d. Prins, R., & H. H. Hobbs, Jr. A new crayfish of the subgenus Puncticambarus from the Savannah River drainage with notes on Cambarus (P.) rebur- rus Prins (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 84:411—420. Cambarus chaugaensis.

1972e. Hobbs, H. H., Jr, & M. R. Cooper. A new troglobitic crayfish from Oklahoma (Decapoda: As-

492

tacidae).—Proceedings of the Biological Society of Washington 85:49—56. Cambarus tartarus.

1972f. Hobbs, H. H., Jr, & E. T. Hall, Jr. A new cray- fish from the Tallapoosa River in Georgia (Decap- oda: Astacidae).—Proceedings of the Biological So- ciety of Washington 85:151—161. Cambarus engli- shi.

1972g. Crayfishes (Astacidae) of North and Middle America.—Biota of Freshwater Ecosystems, Identi- fication Manual No. 9:173 pp. United States Envi- ronmental Protection Agency, Water Pollution Con- trol Research Series.

1973a. Hobbs, H. H., Jr, & R. W. Bouchard. A new crayfish from the Cumberland River system with notes on Cambarus carolinus (Erichson).—Proceed- ings of the Biological Society of Washington 86:41— 68. Cambarus cumberlandensis.

1973b. Three new troglobitic decapod crustaceans from Oaxaca, Mexico. In R. E. Mitchell & J. Red- dell, eds., Studies on the cavernicole fauna of Mex- ico and adjacent regions.—Bulletin, Association for Mexican Cave Studies 5:25—38. Neopalaemon, N. nahuatlus, Procambarus oaxacae oaxacae, P. oax- acae reddelli.

1973c. Hobbs, H. H., Jr, & H. H. Hobbs III. The genus Sphaeromicola (Ostracoda, Entocytheridae) in Mex- ico. In R. E. Mitchell & J. Reddell, eds. Studies on the cavernicole fauna of Mexico and adjacent regions.—Bulletin, Association for Mexican Cave Studies 5:39—42. S. coahuiltecae.

1973d. Two new troglobitic shrimp (Decapoda: Al- pheidae and Palaemonidae) from Oaxaca, Mexico. In R. E. Mitchell & J. Reddell, eds., Studies on the cavernicole fauna of Mexico and adjacent regions.—Bulletin, Association for Mexican Cave Studies 5:73-80. Alpheopsis stygicola, Macrobra- chium villalobosi.

1973e. New species and relationships of the members of the genus Fallicambarus.—Proceedings of the Biological Society of Washington 86:461—482. Creaserinus, F. jeanae, F. spectrum.

1974a. Synopsis of the families and genera of cray- fishes (Crustacea: Decapoda).—Smithsonian Contri- butions to Zoology 164:32 pp.

1974b. Villalobos Figueroa, A., & H. H. Hobbs, Jr. Three new crustaceans from La Media Luna, San Luis Potosi, Mexico.—Smithsonian Contributions to Zoology 174:18 pp. Ankylocythere barbouri, Palae- monetes lindsayi, Procambarus roberti.

1974c. A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cambari- dae).—Smithsonian Contributions to Zoology 166: 161 pp.

1974d. Hobbs, H. H., Jr, & E. T. Hall, Jr. Crayfishes (Decapoda: Astacidae). Pp. 195-214 in C. W. Hart, Jr. & S. L. H. Fuller, eds., Pollution ecology of fresh- water invertebrates. 389 pp. Academic Press, New York.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1975a. Key to the troglobitic crayfishes of Florida. Pp. 14, 15 in K. Relyea & B. Sutton, A new troglobitic crayfish of the genus Procambarus from Florida (Decapoda: Astacidae).—Tulane Studies in Zoology and Botany 19(1&2):8—-16.

1975b. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods from Tennessee with a key to the species of the genus Ascetocythere.—Proceedings of the Biological Society of Washington 88:5—20. A. bouchardi, A. pseudolita, A. triangulata, A. veruta, Dactylocythere crena, D. scissura, Psittocythere, P. psitta.

1975c. New crayfishes (Decapoda: Cambaridae) from the southern United States and Mexico.—Smithson- ian Contributions to Zoology 201:34 pp. Fallicam- barus caesius, F. danielae, Procambarus clemmeri, P. geminus, P. marthae, P. medialis, P. xochitlanae.

1976a. Adaptations and convergence in North Ameri- can crayfishes. Pp. 541-551 in J. W. Avault, Jr., ed., Freshwater crayfish.—Papers from the Second In- ternational Crayfish Symposium, Baton Rouge, Lou- isiana, USA, 1974. 676 pp. Louisiana State Univer- sity.

1976b. Bouchard, R. W., & H. H. Hobbs, Jr A new subgenus and two new species of crayfishes of the genus Cambarus (Decapoda: Cambaridae) from the southern United States—Smithsonian Contributions to Zoology 224:15 pp. C. cracens, C. nodosus, Ex- ilicambarus.

1976c. Hobbs, H. H., Jr., & D. S. Lee. A new troglob- itic crayfish (Decapoda, Cambaridae) from penin- sular Florida.—Proceedings of the Biological Soci- ety of Washington 89:383-—391. Procambarus franzi.

1976d. Hobbs, H. H., Jr, & M. Walton. New entocy- therid ostracods from Kentucky and Tennessee.— Proceedings of the Biological Society of Washing- ton 89:393—404. Ascetocythere riopeli, Dactylocy- there apheles, D. brachydactylus, D. demissa.

1976e. Hobbs Ill, H. H., & H. H. Hobbs, Jr. On the troglobitic shrimps of the Yucatan Peninsula, Mex- ico (Decapoda: Atyidae and Palaemonidae).— Smithsonian Contributions to Zoology 240:23 pp. Typhlatya campecheae, T. mitchelli.

1977a. The crayfish Bouchardina robisoni, a new ge- nus and species (Decapoda, Cambaridae) from southwestern Arkansas.—Proceedings of the Bio- logical Society of Washington 89:733-742.

1977b. Hobbs, H. H., Jr., & D. J. Peters. The entocy- therid ostracods of North Carolina—Smithsonian Contributions to Zoology 247:73 pp. Aphelocythere, A. acuta, Dactylocythere isabelae, D. peedeensis, Donnaldsoncythere leptodrilus, Entocythere costata, Harpagocythere baileyi.

1977c. Hobbs, Jr., H. H., H. H. Hobbs III, & M. A. Daniel. A review of the troglobitic decapod crusta- ceans of the Americas.—Smithsonian Contributions to Zoology 244:183 pp.

1977d. Manning, R. B., & H. H. Hobbs, Jr. Decapoda.

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Pp 157-162 in S. H. Hurlbert, ed., Biota Acuatica de Sudameérica Austral. San Diego State University, San Diego, California, 342 pp.

1977e. A new crayfish (Decapoda: Cambaridae) from San Luis Potosi, Mexico.—Proceedings of the Bio- logical Society of Washington 90(2):412—419. Pro- cambarus strenthi.

1977f. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods of the genus Dactylocythere.—Pro- ceedings of the Biological Society of Washington 90(3):600—614. D. astraphes, D. corvus, D. promi- nula, D. spinescens.

1978. Cave-inhabiting crayfishes of Chiapas, Mexico (Decapoda: Cambaridae). Jn Subterranean Fauna of Mexico, part III, Further results of the Italian zoo- logical missions to Mexico sponsored by the Na- tional Academy of Lincei (1973 and 1975).—Ac- cademia Nazionale dei Lincei, Problemi Attuali di Scienza e di Cultura, Sezione: Missioni ed Esplor- azioni—I [for 1977] 171:197—206. Procambarus sbordonii.

1979a. Hobbs, H. H., Jr., & D. J. Peters. A substitute name for the homonym Aphelocythere Hobbs & Pe- ters (Ostracoda: Entocytheridae)—Proceedings of the Biological Society of Washington 91(4):1037. Waltoncythere.

1979b. A new crayfish from the Ouachita River Basin in Arkansas (Decapoda: Cambaridae).—Proceedings of the Biological Society of Washington 92(4):804— 811. Procambarus reimeri.

1980a. Atya gabonensis (Decapoda Atyidae) in the Western Hemisphere.—Crustaceana 38(1):111, 112.

1980b. New dwarf crayfishes (Decapoda: Cambaridae) from Mexico and Florida.—Proceedings of the Bi- ological Society of Washington 93(1):194-207. Cambarellus blacki, C. chihuahuae.

1980c. Cooper, M. R., & H. H. Hobbs, Jr. New ‘and little-known crayfishes of the virilis section of genus Orconectes (Decapoda: Cambaridae) from the southeastern United States.—Smithsonian Contri- butions to Zoology 320:44 pp. O. chickasawae, O. cooperi, O. holti.

1980d. A new pseudothelphusid crab from the State of Jalisco, Mexico.—Proceedings of the Biological So- ciety of Washington 93(2):357—361. Pseudothelphu- Sa Seiferti.

1981la. Villalobos Figueroa, A., & H. H. Hobbs, Jr. A new dwarf crayfish from the Pacific versant of Mex- ico (Decapoda: Cambaridae).—Proceedings of the Biological Society of Washington 94(2):492—502. Cambarellus prolixus.

1981b. The crayfishes of Georgia.—Smithsonian Con- tributions to Zoology 318:549 pp. Cambarus acan- thura, C. coosae, C. coosawattae, C. fasciatus, C. georgiae, C. harti, C. hiwasseensis, C. manningi, C. parrishi, C. reflexus, C. scotti, C. speciosus, C. stri- gosus, C. truncatus, Distocambarus, Procambarus

493

caritus, P. devexus, P. petersi, P. pubischelae defi- ciens, P. talpoides.

1982a. A new crayfish (Decapoda: Cambaridae) from the State of Puebla, Mexico, with new locality re- cords for Procambarus (Villalobosus) xochitlanae and entocytherid ostracod symbionts.—Association for Mexican Cave Studies Bulletin 8:39—44 (also Texas Memorial Museum Bulletin 28:39—44). Pro- cambarus cuetzalanae.

1982b. Hobbs, H. H., Jr, & A. G. Grubbs. Description of a new troglobitic crayfish from Mexico and a list of Mexican crayfishes reported since the publication of the Villalobos Monograph (1955) (Decapoda, Cambaridae).—Association for Mexican Cave Stud- ies Bulletin 8:45—SO (also Texas Memorial Museum Bulletin 28:45—-50). Procambarus xilitlae.

1982c. Hobbs, H. H., Jr., & D. J. Peters. The entocy- therid ostracod fauna of northern Georgia.—Pro- ceedings of the Biological Society of Washington 95(2):297-3 18.

1982d. Hobbs, H. H., Jr, & H. W. Robison. A new crayfish of the genus Procambarus from southwest- ern Arkansas.—Proceedings of the Biological So- ciety of Washington 95(3):545—-553. P. parasimu- lans.

1982e. Hobbs, H. H., Jr., & C. W. Hart, Jr. The shrimp genus Atya (Decapoda: Atyidae).—Smithsonian Contributions to Zoology 364:143 pp. A. brachyr- hinus.

1982f. On the distribution of the genus Procambarus. Pp. 2, 3 [Abstract] in J. E Payne, ed., Crayfish dis- tribution patterns, American Society of Zoologists.

1983a. Franz, R., & H. H. Hobbs, Jr. Procambarus (Ortmannicus) leitheuseri, new species, another troglobitic crayfish (Decapoda: Cambaridae) from peninsular Florida.—Proceedings of the Biological Society of Washington 96(2):323—332.

1983b. The African shrimp genus Potamalpheops in Mexico (Decapoda: Alpheidae).—Crustaceana 44(2):221—224.

1983c. Hobbs, H. H., Jr., & P H. Carlson. Distocam- barus (Decapoda: Cambaridae) elevated to generic rank, with an account of D. crockeri, new species, from South Carolina.—Proceedings of the Biologi- cal Society of Washington 96(3):420—428.

1983d. Distocambarus (Fitzcambarus) carlsoni, a new subgenus and species of crayfish (Decapoda: Cam- baridae) from South Carolina.—Proceedings of the Biological Society of Washington 96(3):429—439.

1983e. Hobbs, H. H., Jr., & A. C. McClure. On a small collection of entocytherid ostracods with the de- scriptions of three new species.—Proceedings of the Biological Society of Washington 96(4):770-779. Ankylocythere carpenteri, Ascetocythere jezerinaci, Ornithocythere thomai.

1983f. Translation of Cambarinos de la Fauna Mexi- cana (Crustacea Decapoda) [Crayfishes of Mexico Crustacea: Decapoda], Alejandro Villalobos. Smith-

494

sonian Institution Libraries & National Science Foundation. 276 pp. Washington, D.C.

1984a. On the distribution of the crayfish genus Pro- cambarus (Decapoda: Cambaridae).—Journal of Crustacean Biology 4(1):12—24.

1984b. Burr, B. M., & H. H. Hobbs, Jr. Additions to the crayfish fauna of Kentucky, with new locality records for Cambarellus shufeldtii.—Transactions of the Kentucky Academy of Sciences 45(1—2):14—18.

1985a. Hobbs, H. H., Jr., & P H. Carlson. A new mem- ber of the genus Distocambarus (Decapoda: Cam- baridae) from the Saluda Basin, South Carolina.— Proceedings of the Biological Society of Washing- ton 98(1):81—89. D. youngineri.

1985b. Hobbs, H. H., Jr, & H. W. Robison. A new burrowing crayfish (Decapoda: Cambaridae) from southwestern Arkansas.—Proceedings of the Bio- logical Society of Washington 98(4):1035—1041. Fallicambarus harpi.

1986a. Andolshek, M. D., & H. H. Hobbs, Jr. The entocytherid ostracod fauna of southeastern Geor- gia.—Smithsonian Contributions to Zoology 424:43 pp. Ankylocythere spargosis, Entocythere prisma.

1986b. Hobbs, H. H., Jr, & R. Franz. New troglobitic crayfish with comments on its relationship to epi- gean and other hypogean crayfishes of Florida. Journal of Crustacean Biology 6(3):509-—519. Pro- cambarus delicatus.

1986c. Hobbs, H. H., Jr, & A. G. Grubbs. Notes on the crayfish Procambarus (Ortmannicus) xilitlae (Decapoda: Cambaridae).—Proceedings of the Bio- logical Society of Washington 99(4):735-738.

1986d. Highlights of a half century of crayfishing. Pp. 12—23 in P. Brinck, ed., Freshwater Crayfish VI, Pa- pers from the Sixth International Symposium of As- tacology, Lund Sweden, 13-15 August 1984. 281 pp. International Association of Astacology.

1986e. A new troglobitic crab (Crustacea: Decapoda: Pseudothelphusidae) from Belize. Pp. 1—4 in J. R. Reddell, ed., Studies on the cave and endogean fau- na of North America.—Texas Memorial Museum Speleological Monographs 1:1—4. Typhlopseudoth- elphusa acanthochela.

1987a. On the identity of Astacus (Cambarus) mexi- canus Erichson (1846) and Cambarus aztecus Saus- sure (1857) (Decapoda: Cambaridae) with the de- scription of Procambarus olmecorum, new species, from Veracruz, Mexico.—Proceedings of the Bio- logical Society of Washington 100(1):198—215.

1987b. A review of the crayfish genus Astacoides (De- capoda: Parastacidae).—Smithsonian Contributions to Zoology 443:50 pp. A. crosnieri, A. petiti.

1987c. Hobbs, H. H., Jr, & M. Whiteman. A new, economically important crayfish (Decapoda: Cam- baridae) from the Neches River Basin, Texas, with a key to the subgenus Fallicambarus.—Proceedings of the Biological Society of Washington 100(2): 403-411. Fallicambarus devastator.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1987d. Hobbs, H. H., Jr., & A. V. Brown. A new trog- lobitic crayfish from northwestern Arkansas (Decap- oda: Cambaridae).—Proceedings of the Biological Society of Washington 100(4):1040—1048. Camba- rus aculabrum.

1988a. Crayfish distribution, adaptive radiation and evolution. Pp. 52-82 in D. M. Holdich & R. S. Low- ery, eds., Freshwater crayfish: biology, management and exploitation. 498 pp. Croom Helm: London and Sydney; Timber Press: Portland, Oregon.

1988b. Hobbs, H. H., Jr., & H. W. Robison. The cray- fish subgenus Girardiella (Decapoda: Cambaridae) in Arkansas, with the descriptions of two new spe- cies and a key to the members of the Gracilis group in the genus Procambarus.—Proceedings of the Bi- ological Society of Washington 101(2):391—413. Procambarus ferrugineus, P. regalis.

1988c. Hobbs, H. H., Jr, & W. L. Pflieger. Cambarus (Erebicambarus) maculatus, new crayfish (Decapo- da: Cambaridae) from the Meramec River Basin of Missouri.—Proceedings of the Biological Society of Washington 101(3):644—-652.

1989a. Burrowing crayfishes in Virginia. The Virginia Explorer 5(2):5-7.

1989b. Hobbs, H. H., Jr., & D. J. Peters. New records of entocytherid ostracods infesting burrowing cray- fishes, with the description of a new species, Asce- tocythere stockeri.—Proceedings of the Biological Society of Washington 102(2):324—330.

1989c. Rodriguez, G., & H. H. Hobbs, Jr. Freshwater crabs associated with caves in southern Mexico and Belize, with descriptions of three new species (Crus- tacea: Decapoda).—Proceedings of the Biological Society of Washington 102(2):394—400. Odonto- thelphusa monodontis, Potamocarcinus leptomelus, Typhlopseudothelphusa hyba.

1989d. Rodriguez, G. & H. H. Hobbs, Jr. A new cav- ernicolous crab, Zilchia falcata, from Guatemala, with notes on the genera of the Potamocarcinini (Crustacea Decapoda, Pseudothelphusidae).—Bul- letin du Muséum National d’ Histoire Naturelle, Par- is, series 4, 11(section A) (1):183—192.

1989e. Hobbs, H. H. Jr., & H. H. Hobbs III. New lo- cality records for two poorly known Mexican fresh- water shrimps (Decapoda, Palaemonidae).—Crusta- ceana 57(2):220—222.

1989f. Hobbs, Jr., H. H., & H. W. Robison. On the crayfish genus Fallicambarus (Decapoda: Cambar- idae) in Arkansas, with notes on the Fodiens com- plex and descriptions of two new species.—Pro- ceedings of the Biological Society of Washington 102(3):651-697. F. gilpini, F. petilicarpus.

1989g. An illustrated checklist of the American cray- fishes (Decapoda: Astacidae, Cambaridae, and Par- astacidae).—Smithsonian Contributions to Zoology 480:236 pp.

1990a. On the crayfishes (Decapoda: Cambaridae) of the Neches River Basin of eastern Texas with the

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descriptions of three new species.—Proceedings of the Biological Society of Washington 103(3):573— 597. Procambarus kensleyi, P. nechesae, P. nigro- cinctus.

1990b. Hobbs, H. H., Jr., & H. H. Hobbs III. A new crayfish (Decapoda: Cambaridae) from southeastern Texas.—Proceedings of the Biological Society of Washington 103(3):608-613. Procambarus zonan- gulus.

1991a. Hobbs, H. H., Jr, & M. Whiteman. Notes on the burrows, behavior, and color of the crayfish Fal- licambarus (F.) devastator (Decapoda: Cambari- dae).—The Southwestern Naturalist 36(1):127—135.

1991b. Hobbs, H. H., Jr, & R. Franz. A new troglob- itic crayfish, Procambarus (Lonnbergius) morrisi, (Decapoda: Cambaridae) from Florida.—Proceed- ings of the Biological Society of Washington 104(1):55-63.

1991c. Hobbs, H. H., Jr, & D. J. Peters. Additional records of entocytherid ostracods infesting burrow- ing crayfishes, with the description of five new spe- cies.—Proceedings of the Biological Society of Washington 104(1):64—75. Ankylocythere prolata, Dactylocythere guyandottae, D. lepta, D. pygidion, Plectocythere kentuckiensis.

1991d. Hobbs, H. H., Jr, & H. H. Hobbs III. An il- lustrated key to the crayfishes of Florida (based on first form males).—Florida Scientist 54(1):13—24.

199le. Procambarus (Girardiella) steigmani, a new crayfish (Decapoda, Cambaridae) from a long-grass prairie in northeastern Texas.—Proceedings of the Biological Society of Washington 104(2):309-316.

1991f. A new generic assignment for a South Ameri- can crayfish (Decapoda: Parastacidae) with revised diagnoses of the South American genera and com- ments on the parastacid mandible.—Proceedings of the Biological Society of Washington 104(4):800— 811. Virilastacus.

1992. Hobbs, H. H., Jr., & R. Franz. Procambarus (Ortmannicus) attiguus, a new troglobitic crayfish (Decapoda, Cambaridae) from the Saint Johns River Basin, Florida.—Proceedings of the Biological So- ciety of Washington 105(2):359-—365.

1993. Hobbs, H. H., Jr, & D. J. Peters. New records of entocytherid ostracods infesting burrowing and cave-dwelling crayfishes, with descriptions of two new species.—Proceedings of the Biological Soci- ety of Washington 106(3):455—466. Dactylocythere cryptoteresis, Phymocythere lophota.

1994. Hobbs, H. H., Jr., & R. W. Bouchard. Cambarus (Cambarus) angularis, a new crayfish (Decapoda: Cambaridae) from the Tennessee River Basin of northeastern Tennessee and Virginia.—Jeffersoniana 5:1-13.

1995a. Hobbs, H. H., Jr., & H. H. Hobbs III. Macro- brachium catonium, a new troglobitic shrimp from the Cayo District of Belize (Crustacea: Decapoda:

495

Palaemonidae).—Proceedings of the Biological So-

ciety of Washington 108(1):50-—53. 1995b. Hobbs, H. H., Jr., & H. H. Hobbs III. Procam-

barus (Ortmannicus) nueces (Decapoda: Cambari- dae), a new crayfish from the Nueces River Basin,

Texas.—Proceedings of the Biological Society of

Washington 108(1):54—60.

At the time of his death, Hobbs was com- pleting a manuscript entitled “‘A compara- tive study of functional morphology of the male reproductive systems in the Astacidea (Crustacea: Decapoda) with emphasis on the freshwater crayfishes.”’ Hobbs III plans to complete preparing this manuscript for publication.

Taxa Named by Horton H. Hobbs, Jr.

The family, genera and subgenera, spe- cies and subspecies named by Hobbs and colleagues are listed alphabetically and are cross-referenced to Hobbs’s bibliography, above. We provide the repository and cat- alogue number for all holotypes of species and subspecies. Most of the holotypes of taxa named by Hobbs are in the collections of the National Museum of Natural History, Smithsonian Institution, Washington (USNM). One holotype is in The Natural History Museum, London (BMNH) and several are in the Muséum National d’ Histoire Naturelle, Paris (MNHN) as well as The Academy of Natural Sciences of Philadelphia (ANSP). When a holotype has been deposited in a museum other than the USNM, catalogue numbers of USNM para- types are provided.

Acucauda Hobbs, 1972a.

Alpheopsis stygicola Hobbs, 1973d. Holo- type USNM 143629.

Ankylocythere barbouri Villalobos Figu- eroa & Hobbs, 1974b. Holotype USNM 149159.

Ankylocythere carpenteri Hobbs & Mc- Clure, 1983e. Holotype USNM 204402.

Ankylocythere harmani Hobbs, 1966f. Ho- lotype USNM 123532.

Ankylocythere hyba Hobbs & Walton, 1963a. Holotype USNM 108016.

496

Ankylocythere maya Hobbs, 1971b. Holo- type USNM 128822. Ankylocythere prolata Hobbs & Peters, 1991c. Holotype USNM 235511. Ankylocythere spargosis Andolshek & Hobbs, 1986. Holotype USNM 213651. Ankylocythere toltecae Hobbs, 1971b. Ho- lotype USNM 128823. Ankylocythere villalobosi Hobbs, Holotype USNM 128825. Aphelocythere Hobbs & Peters, 1977b. Aphelocythere acuta Hobbs & Peters, 1977b. Holotype USNM 155324. Ascetocythere batchi Hobbs & Walton, 1968d. Holotype USNM 123321. Ascetocythere bouchardi Hobbs & Walton, 1975b. Holotype USNM 150640. Ascetocythere coryphodes Hobbs & Hart, 1966d. Holotype USNM 113449. Ascetocythere cosmeta Hobbs & Hart, 1966d. Holotype USNM 113453. Ascetocythere didactylata Hobbs & Hart, 1966d. Holotype USNM 113448. Ascetocythere hoffmani Hobbs & Hart, 1966d. Holotype USNM 113441. Ascetocythere holti Hobbs & Walton, 1970b. Holotype USNM 126974. Ascetocythere hyperoche Hobbs & Hart, 1966d. Holotype USNM 113444. Ascetocythere jezerinaci Hobbs & Mc- Clure, 1983e. Holotype USNM 204400. Ascetocythere lita Hobbs & Hobbs III, 1970c. Holotype USNM 126251. Ascetocythere myxoides Hobbs & Hart, 1966d. Holotype USNM 113451. Ascetocythere ozalea Hobbs & Hart, 1966d. Holotype USNM 113442. Ascetocythere pseudolita Hobbs & Walton, 1975b. Holotype USNM 150642. Ascetocythere riopeli Hobbs & Walton, 1976d. Holotype USNM 155317. Ascetocythere sclera Hobbs & Hart, 1966d. Holotype USNM 113445. Ascetocythere stockeri Hobbs & Peters, 1989b. Holotype USNM 240114. Ascetocythere triangulata Hobbs & Walton, 1975b. Holotype USNM 150643. Ascetocythere veruta Hobbs & Walton, 1975b. Holotype USNM 150645.

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Astacoides crosnieri Hobbs, 1987b. Holo- type MNHN As328; paratype USNM NST.

Astacoides petiti Hobbs, 1987b. Holotype MNHN As211; paratype USNM 218799.

Atya brachyrhinus Hobbs & Hart, 1982e. Holotype BMNH 1972:539; paratype USNM 184857.

Austrocambarus Hobbs, 1972a.

Aviticambarus Hobbs, 1969c.

Barbicambarus Hobbs, 1969c.

Bouchardina Hobbs, 1977a.

Bouchardina robisoni Hobbs, 1977a. Ho- lotype USNM 147146.

Cambarellus blacki Hobbs, 1980b. Holo- type USNM 148901.

Cambarellus chihuahuae Hobbs, Holotype USNM 148895.

Cambarellus diminutus Hobbs, 1945c. Ho- lotype USNM 81554.

Cambarellus ninae Hobbs, 1950a. Holotype USNM 89768.

Cambarellus prolixus Villalobos-Figueroa & Hobbs, 198la. Holotype USNM 177206.

Cambarellus puer Hobbs, 1945c. Holotype USNM 81556.

Cambarellus schmitti Hobbs, 1942d. Ho- lotype USNM 81291.

Cambaridae Hobbs, 1942d.

Cambarus acanthura Hobbs, 1981b. Holo- type USNM 129758.

Cambarus aculabrum Hobbs & Brown, 1987d. Holotype USNM 219149.

Cambarus angularis Hobbs & Bouchard, 1994. Holotype USNM 260252.

Cambarus blandingii cuevachicae Hobbs, 1941a. Holotype USNM 80030.

Cambarus bouchardi Hobbs, 1970e. Holo- type USNM 130295.

Cambarus brachydactylus Hobbs, 1953a. Holotype USNM 93155.

Cambarus byersi Hobbs, 1941b. Holotype USNM 79342.

Cambarus catagius Hobbs & Perkins, 1967c. Holotype USNM 117799.

Cambarus chaugaensis Prins & Hobbs, 1972d. Holotype USNM 131926.

1980b.

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Cambarus conasaugaensis Hobbs & Hobbs III, 1962a. Holotype USNM 107156.

Cambarus coosae Hobbs, 1981b. Holotype USNM 145603.

Cambarus coosawattae Hobbs, 1981b. Ho- lotype USNM 148112.

Cambarus cracens Bouchard & Hobbs, 1976b. Holotype USNM 146082.

Cambarus cristatus Hobbs, 1955a. Holo- type USNM 96985.

Cambarus cryptodytes Hobbs, 1941b. Ho- lotype USNM 79339.

Cambarus cumberlandensis Hobbs & Bou- chard, 1973a. Holotype USNM 132989.

Cambarus cymatilis Hobbs, 1970e. Holo- type USNM 129860.

Cambarus englishi Hobbs & Hall, 1972f. Holotype USNM 131700.

Cambarus fasciatus Hobbs, 1981b. Holo- type USNM 147917.

Cambarus floridanus Hobbs, 1941b. Holo- type USNM 79341.

Cambarus friaufi Hobbs, 1953a. Holotype USNM 93157.

Cambarus gentryi Hobbs, 1970d. Holotype USNM 130283.

Cambarus georgiae Hobbs, 1981b. Holo- type USNM 118944.

Cambarus halli Hobbs, 1968a. Holotype USNM 129288.

Cambarus harti Hobbs, 1981b. Holotype USNM 148348.

Cambarus hedgpethi Hobbs, 1948f. Holo- type USNM 85146.

Cambarus hiwasseensis Hobbs, 1981b. Ho- lotype USNM 129366.

Cambarus howardi Hobbs & Hall, 1969d. Holotype USNM 129866.

Cambarus hubrichti Hobbs, 1952d. Holo- type USNM 92295.

Cambarus hubbelli Hobbs, 1940a. Holo- type USNM 76593.

Cambarus jonesi Hobbs & Barr, 1960b. Holotype USNM 104407.

Cambarus kilbyi Hobbs, 1940a. Holotype USNM 76594.

Cambarus lucifugus alachua Hobbs, 1940a. Holotype USNM 76592.

497

Cambarus lucifugus lucifugus Hobbs, 1940a. Holotype USNM 77916.

Cambarus lunzi Hobbs, 1940b. Holotype USNM 79301.

Cambarus maculatus Hobbs & Pflieger, 1988c. Holotype USNM 219292.

Cambarus manningi Hobbs, 1981b. Holo- type USNM 147911.

Cambarus nerterius Hobbs, 1964c. Holo- type USNM 111295.

Cambarus nodosus Bouchard & Hobbs, 1976b. Holotype USNM 146756.

Cambarus obeyensis Hobbs & Shoup, 1947b. Holotype USNM 82260.

Cambarus pallidus Hobbs, 1940a. Holotype USNM 76591.

Cambarus parrishi Hobbs, 1981b. Holo- type USNM 144957.

Cambarus parvoculus Hobbs & Shoup, 1947b. Holotype USNM 82259.

Cambarus pictus Hobbs, 1940a. Holotype USNM 76596.

Cambarus pristinus Hobbs, 1965b. Holo- type USNM 115528.

Cambarus prominens Hobbs, 1966b. Ho- lotype USNM 115603.

Cambarus rathbunae Hobbs, 1940a. Holo- type USNM 76595.

Cambarus reduncus Hobbs, 1956a. Holo- type USNM 99183.

Cambarus reflexus Hobbs, 1981b. Holotype USNM 148116.

Cambarus rogersi Hobbs, 1938b. Holotype USNM 75120.

Cambarus scotti Hobbs, 1981b. Holotype USNM 146479.

Cambarus speciosus Hobbs, 1981b. Holo- type USNM 146023.

Cambarus sphenoides Hobbs, 1968a. Ho- lotype USNM 129325.

Cambarus spicatus Hobbs, 1956b. Holo- type USNM 99323.

Cambarus strigosus Hobbs, 1981b. Holo- type USNM 148284.

Cambarus tartarus Hobbs & Cooper, 1972e. Holotype USNM 131951.

Cambarus truncatus Hobbs, 1981b. Holo- type USNM 116966.

498

Cambarus unestami Hobbs & Hall, 1969d. Holotype USNM 129863. Cambarus zophonastes Hobbs & Bedinger, 1964b. Holotype USNM 108356. Capillicambarus Hobbs, 1972a. Creaserinus Hobbs, 1973e. Cymocythere gonia Hobbs & Hart, 1966d. Holotype USNM 113455. Dactylocythere apheles Hobbs & Walton, 1976d. Holotype USNM 155319. Dactylocythere astraphes Hobbs & Walton, 1977f. Holotype USNM 169073. Dactylocythere brachydactylus Hobbs & Walton, 1976d. Holotype USNM 155321. Dactylocythere brachystrix Hobbs & Wal- ton, 1966e. Holotype USNM 111251. Dactylocythere coloholca Hobbs & Hobbs III, 1970c. Holotype USNM 126253. Dactylocythere cooperorum Hobbs & Wal- ton, 1968d. Holotype USNM 123322. Dactylocythere corvus Hobbs & Walton, 1977f. Holotype USNM 169075. Dactylocythere crena Hobbs & Walton, 1975b. Holotype USNM 150646. Dactylocythere cryptoteresis Hobbs & Pe- ters, 1993. Holotype USNM 260073. Dactylocythere demissa Hobbs & Walton, 1976d. Holotype USNM 155323. Dactylocythere enoploholea Hobbs & Wal- ton, 1970b. Holotype USNM 126973. Dactylocythere guyandottae Hobbs & Pe- ters, 1991c. Holotype USNM 235513. Dactylocythere isabelae Hobbs & Peters, 1977b. Holotype USNM 155326. Dactylocythere jeanae Hobbs, 1967a. Ho- lotype USNM 113475. Dactylocythere lepta Hobbs & Peters, 1991c. Holotype USNM 235514. Dactylocythere macroholca Hobbs & Hobbs III, 1970c. Holotype USNM 126255. Dactylocythere myura Hobbs & Walton, 1970b. Holotype USNM 126975. Dactylocythere pachysphyrata Hobbs & Walton, 1966e. Holotype USNM 111253. Dactylocythere peedeensis Hobbs & Peters, 1977b. Holotype USNM 155328. Dactylocythere phoxa Hobbs, 1967a. Ho- lotype USNM 113477.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Dactylocythere prinsi Hobbs & Walton, 1968d. Holotype USNM 123325. Dactylocythere prominula Hobbs & Wal- ton, 1977f. Holotype USNM 169077. Dactylocythere pughae Hobbs & Hobbs III, 1970c. Holotype USNM 126257. Dactylocythere pygidion Hobbs & Peters, 1991c. Holotype USNM 235516. Dactylocythere scissura Hobbs & Walton, 1975b. Holotype USNM 150648. Dactylocythere spinata Hobbs & Walton, 1970b. Holotype USNM 126972. Dactylocythere spinescens Hobbs & Wal- ton, 1977f. Holotype USNM 169079. Dactylocythere xystroides Hobbs & Walton, 1963a. Holotype USNM 108029. Depressicambarus Hobbs, 1969c. Distocambarus Hobbs, 1981b. Distocambarus carlsoni Hobbs, 1983d. Ho- lotype USNM 178599. Distocambarus crockeri Hobbs & Carlson, 1983c. Holotype USNM 178582. Distocambarus youngineri Hobbs & Carl- son, 1985a. Holotype USNM 208414. Donnaldsoncythere ardis Hobbs & Walton, 1963b. Holotype USNM 108018. Donnaldsoncythere cayugaensis Hobbs & Walton, 1966e. Holotype USNM 111255. Donnaldsoncythere ileata Hobbs & Walton, 1963b. Holotype USNM 108027. Donnaldsoncythere leptodrilus Hobbs & Peters, 1977b. Holotype USNM 155330. Donnaldsoncythere scalis Hobbs & Walton, 1963b. Holotype USNM 108024. Donnaldsoncythere truncata Hobbs & Wal- ton, 1963b. Holotype USNM 108021. Entocythere ambophora Walton & Hobbs, 1959c. Holotype USNM 105953. Entocythere arcuata Hart & Hobbs, 1961b. Holotype ANSP 6233. Entocythere asceta Hobbs & Walton, 1962b. Holotype USNM 108174. Entocythere barri Hart & Hobbs, 1961b. Holotype ANSP 6231. Entocythere chalaza Hobbs & Walton, 1962b. Holotype USNM 108179. Entocythere costata Hobbs & Peters, 1977b. Holotype USNM 155334.

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Entocythere cyma Hobbs & Walton, 1960a. Holotype USNM 105962. Entocythere daphnioides Hobbs, 1955b. Holotype USNM 98406. Entocythere falcata Hobbs & Walton, 1961la. Holotype USNM 105966. Entocythere hamata Hobbs, 1957a. Holo- type USNM 100938. Entocythere hiwasseensis Hobbs & Walton, 1961a. Holotype USNM 105963. Entocythere kanawhaensis Hobbs & Wal- ton, 1966e. Holotype USNM 111257. Entocythere lucifuga Walton & Hobbs, 1959c. Holotype USNM 105957. Entocythere mecoscapha Hobbs & Walton, 1960a. Holotype USNM 105960. Entocythere pholetera Hart & Hobbs, 1961b. Holotype ANSP 6238. Entocythere phyma Hobbs & Walton, 1962b. Holotype USNM 108178. Entocythere prionata Hart & Hobbs, 1961b. Holotype ANSP 6235. Entocythere prisma Andolshek & Hobbs, 1986a. Holotype USNM 213649. Entocythere reddelli Hobbs & Walton, 1968d. Holotype USNM 123330. Entocythere runki Hobbs, 1955b. Holotype USNM 98410. Entocythere simondsi Hobbs & Walton, 1960a. Holotype USNM 105956. Entocythere steevesi Hart & Hobbs, 1961b. Holotype ANSP 6236. | Entocythere tuberosa Hart & Hobbs, 1961b. Holotype ANSP 6340. Entocythere tyttha Hobbs & Hobbs III, 1970c. Holotype USNM 126258. Entocythere ungulata Hart & Hobbs, 1961b. Holotype ANSP 6234. Entocythere xania Hart & Hobbs, 1961b. Holotype ANSP 6237. Erebicambarus Hobbs, 1969a. Exilicambarus Bouchard & Hobbs, 1976d. Fallicambarus Hobbs, 1969c. Fallicambarus caesius Hobbs, 1975c. Ho- lotype USNM 144921. Fallicambarus danielae Hobbs, 1975c. Ho- lotype USNM 145997. Fallicambarus devastator Hobbs & White- man, 1987c. Holotype USNM 218546.

499

Fallicambarus gilpini Hobbs & Robison, 1989f. Holotype USNM 219511.

Fallicambarus harpi Hobbs & Robison, 1985b. Holotype USNM 217946.

Fallicambarus hortoni Hobbs & Fitzpat- rick, 1970a. Holotype USNM 129895.

Fallicambarus jeanae Hobbs, 1973e. Ho- lotype USNM 144672.

Fallicambarus petilicarpus Hobbs & Rob- ison, 1989f. Holotype USNM 219507. Fallicambarus spectrum Hobbs, 1973e. Ho-

lotype USNM 144674. Fitzcambarus Hobbs, 1983d. Geocythere nessoides Hobbs & Hobbs III, 1970c. Holotype USNM 126259. Hagenides Hobbs, 1972a. Harpagocythere baileyi Hobbs & Peters, 1977b. Holotype USNM 155332. Harpagocythere tertius Hobbs & Walton, 1968d. Holotype USNM 123329. Hiaticambarus Hobbs, 1969c. Jugicambarus Hobbs, 1969c. Lacunicambarus Hobbs, 1969c. Leconticambarus Hobbs, 1972a. Litocythere Hobbs & Walton, 1968d. Litocythere lucileae Hobbs & Walton, 1968d. Holotype USNM 123327. Lonnbergius Hobbs, 1972a. Lordocythere Hobbs & Hobbs III, 1970c. Lordocythere petersi Hobbs & Hobbs III, 1970c. Holotype USNM 126262. Macrobrachium catonium Hobbs & Hobbs III, 1995a. Holotype USNM 260328. Macrobrachium villalobosi Hobbs, 1973d. Holotype USNM 143633. Mexicambarus Hobbs, 1972a. Neopalaemon Hobbs, 1973b. Neopalaemon nahuatlus Hobbs, 1973b. Holotype USNM 143120. Odontothelphusa monodontis Rodriguez & Hobbs, 1989c. Holotype USNM 230078. Orconectes chickasawae Cooper & Hobbs, 1980c. Holotype USNM 144926. Orconectes cooperi Cooper & Hobbs, 1980c. Holotype USNM 147722. Orconectes holti Cooper & Hobbs, 1980c. Holotype USNM 147149. Orconectes incomptus Hobbs & Barr, 1972b. Holotype USNM 130299.

500

Orconectes leptogonopodus Hobbs, 1948b. Holotype USNM 82262.

Orconectes marchandi Hobbs, 1948b. Ho- lotype USNM 82072.

Orconectes propinquus erismophorous Hobbs & Fitzpatrick, 1962d. Holotype USNM 107597.

Orconectes rhoadesi Hobbs, 1949b. Holo- type USNM 87953.

Orconectes shoupi Hobbs, 1948a. Holotype USNM 84072.

Orconectes virginiensis Hobbs, 1951a. Ho- lotype USNM 91659.

Orconectes wrighti Hobbs, 1948e. Holo- type USNM 85144.

Ornithocythere Hobbs, 1967a.

Ornithocythere thomai Hobbs & McClure, 1983e. Holotype USNM 204405.

Ornithocythere waltonae Hobbs, 1967a. Holotype USNM 113472.

Palaemonetes lindsayi Villalobos Figueroa & Hobbs, 1974b. Holotype USNM 149161.

Pennides Hobbs, 1972a.

Phymocythere Hobbs & Hart, 1966d.

Phymocythere lophota Hobbs & Peters, 1993. Holotype USNM 260074.

Plectocythere johnsonae Hobbs & Hart, 1966d. Holotype USNM 113454.

Plectocythere kentuckiensis Hobbs & Pe- ters, 1991c. Holotype USNM 235517.

Potamocarcinus leptomelus Rodriguez & Hobbs, 1989c. Holotype USNM 230080.

Procambarus ancylus Hobbs, 1958b. Ho- lotype USNM 115050.

Procambarus apalachicolae Hobbs, 1942d. Holotype USNM 81272.

Procambarus attiguus Hobbs & Franz, 1992. Holotype USNM 220683.

Procambarus bivittatus Hobbs, 1942d. Ho- lotype USNM 81274.

Procambarus capillatus Hobbs, 1971c. Ho- lotype USNM 131454.

Procambarus caritus Hobbs, 1981b. Holo- type USNM 117598.

Procambarus chacei Hobbs, 1958c. Holo- type USNM 101289.

Procambarus clemmeri Hobbs, 1975c. Ho- lotype USNM 145607.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Procambarus cuetzalanae Hobbs, 1982a. Holotype USNM 177202.

Procambarus delicatus Hobbs & Franz, 1986b. Holotype USNM 218528.

Procambarus devexus Hobbs, 1981b. Ho- lotype USNM 148569.

Procambarus echinatus Hobbs, 1956c. Ho- lotype USNM 99180.

Procambarus econfinae Hobbs, 1942d. Ho- lotype USNM 81276.

Procambarus elegans Hobbs, 1969e. Ho- lotype USNM 129892.

Procambarus enoplosternum Hobbs, 1947a. Holotype USNM 82263.

Procambarus epicyrtus Hobbs, 1958c. Ho- lotype USNM 101286.

Procambarus escambiensis Hobbs, 1942d. Holotype USNM 81278. Procambarus ferrugineus Hobbs & Robi- son, 1988b. Holotype USNM 218841. Procambarus fitzpatricki Hobbs, 1971a. Holotype USNM 131205.

Procambarus franzi Hobbs & Lee, 1976c. Holotype USNM 146992.

Procambarus geminus Hobbs, 1975c. Ho- lotype USNM 145756.

Procambarus geodytes Hobbs, 1942d. Ho- lotype USNM 82263.

Procambarus gibbus Hobbs, 1969e. Holo- type USNM 129804.

Procambarus hirsutus Hobbs, 1958b. Ho- lotype USNM 101148.

Procambarus horsti Hobbs & Means, 1972c. Holotype USNM 132043.

Procambarus howellae Hobbs, 1952c. Ho- lotype USNM 93158.

Procambarus hybus Hobbs & Walton, 1957b. Holotype USNM 99581.

Procambarus jaculus Hobbs & Walton, 1957b. Holotype USNM 99584.

Procambarus kensleyi Hobbs, 1990a. Ho- lotype USNM 219772.

Procambarus latipleurum Hobbs, 1942d. Holotype USNM 81281.

Procambarus leitheuseri Franz & Hobbs, 1983a. Holotype USNM 178361.

Procambarus leonensis Hobbs, 1942d. Ho- lotype USNM 81091.

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Procambarus lepidodactylus Hobbs, 1947c. Holotype USNM 84198.

Procambarus lewisi Hobbs & Walton, 1959a. Holotype USNM 102467.

Procambarus litosternum Hobbs, Holotype USNM 82261.

Procambarus lophotus Hobbs & Walton, 1960c. Holotype USNM 104404.

Procambarus lylei Fitzpatrick & Hobbs, 1971d. Holotype USNM 131533.

Procambarus mancus Hobbs & Walton, 1957b. Holotype USNM 99578.

Procambarus marthae Hobbs, 1975c. Ho- lotype USNM 145994.

Procambarus medialis Hobbs, 1975c. Ho- lotype USNM 144942.

Procambarus milleri Hobbs, 1971f. Holo- type USNM 131257.

Procambarus morrisi Hobbs & Franz, 1991b. Holotype USNM 220374.

Procambarus nechesae Hobbs, 1990a. Ho- lotype USNM 219733.

Procambarus nigrocinctus Hobbs, 1990a. Holotype USNM 219426.

Procambarus niveus Hobbs & Villalobos, 1964a. Holotype USNM 109076.

Procambarus nueces Hobbs & Hobbs III, 1995b. Holotype USNM 260326.

Procambarus oaxacae oaxacae Hobbs, 1973b. Holotype USNM 144341.

Procambarus oaxacae reddelli Hobbs, 1973b. Holotype USNM 144346.

Procambarus okaloosae Hobbs, 1942d. Ho- lotype USNM 81282.

Procambarus olmecorum Hobbs, Holotype USNM 217626.

Procambarus orcinus Hobbs & Means, 1972c. Holotype USNM 132031.

Procambarus parasimulans Hobbs & Rob- ison, 1982d. Holotype USNM 177698.

Procambarus pearsei plumimanus Hobbs & Walton, 1958d. Holotype USNM 100940.

Procambarus pecki Hobbs, 1967e. Holo- type USNM 117684.

Procambarus penni Hobbs, 1951b. Holo- type USNM 91662.

Procambarus petersi Hobbs, 1981b. Holo- type USNM 144960.

1947a.

1987a.

501

Procambarus pubischelae Hobbs, 1942d. Holotype USNM 81284.

Procambarus pubischelae deficiens Hobbs, 1981b. Holotype USNM 133315.

Procambarus pycnogonopodus Hobbs, 1942d. Holotype USNM 81092.

Procambarus pygmaeus Hobbs, 1942d. Ho- lotype USNM 81285.

Procambarus raneyi Hobbs, 1953e. Holo- type USNM 95124.

Procambarus regalis Hobbs & Robison, 1988b. Holotype USNM 219244.

Procambarus reimeri Hobbs, 1979b. Ho- lotype USNM 148880.

Procambarus roberti Villalobos Figueroa & Hobbs, 1974b. Holotype USNM 132345.

Procambarus rodriguezi Hobbs, 1943c. Holotype USNM 81302.

Procambarus rogersi campestris Hobbs, 1942d. Holotype USNM 79384.

Procambarus rogersi expletus Hobbs & Hart, 1959b. Holotype USNM 102125.

Procambarus rogersi ochlocknensis Hobbs, 1942d. Holotype USNM 79383.

Procambarus sbordonii Hobbs, 1978. Ho- lotype USNM 146588.

Procambarus seminolae Hobbs, 1942d. Ho- lotype USNM 81286.

Procambarus shermani Hobbs, 1942d. Ho- lotype USNM 81287.

Procambarus steigmani Hobbs, 1991e. Ho- lotype USNM 220525.

Procambarus strenthi Hobbs, 1977e. Ho- lotype USNM 147725.

Procambarus suttkusi Hobbs, 1953d. Ho- lotype USNM 93730.

Procambarus talpoides Hobbs, 1981b. Ho- lotype USNM 115002.

Procambarus tenuis Hobbs, 1950b. Holo- type USNM 90390.

Procambarus texanus Hobbs, 1971c. Ho- lotype USNM 131457.

Procambarus toltecae Hobbs, 1943c. Ho- lotype USNM 81301.

Procambarus truculentus Hobbs, Holotype USNM 95670.

Procambarus verrucosus Hobbs, 1952a. Holotype USNM 118675.

1954b.

502

Procambarus villalobosi Hobbs, 1969b. Holotype USNM 118644.

Procambarus xilitlae Hobbs & Grubbs, 1982b. Holotype USNM 177140.

Procambarus xochitlanae Hobbs, Holotype USNM 145610.

Procambarus youngi Hobbs, 1942d. Holo- type USNM 81289. Procambarus zonangulus Hobbs & Hobbs III, 1990b. Holotype USNM 220297. Pseudothelphusa seiferti Hobbs, 1980d. Holotype USNM 173884.

Psittocythere Hobbs & Walton, 1975b.

Psittocythere psitta Hobbs & Walton, 1975b. Holotype USNM 150650.

Puncticambarus Hobbs, 1969c.

Remoticambarus Hobbs, 1972a.

Scapulicambarus Hobbs, 1972a.

Sphaeromicola coahuiltecae Hobbs & Hobbs, III, 1973c. Holotype USNM 143122.

Tenuicambarus Hobbs, 1972a.

Thermastrocythere Hobbs 1966e.

Thermastrocythere harti Hobbs & Walton, 1966e. Holotype USNM 111258.

Troglocambarus Hobbs, 1942b.

Troglocambarus maclanei Hobbs, 1942b. Holotype USNM 79385.

Typhlatya campecheae Hobbs III & Hobbs, 1976e. Holotype USNM 151903.

Typhlatya mitchelli Hobbs II & Hobbs, 1976e. Holotype USNM 151904.

Typhlopseudothelphusa acanthochela Hobbs, 1986e. Holotype USNM 216239.

Typhlopseudothelphusa hyba Rodriguez & Hobbs, 1989c. Holotype USNM 150633.

Uncinocythere stubbsi Hobbs & Walton, 1966e. Holotype USNM 111260.

Uncinocythere warreni Hobbs & Walton, 1968d. Holotype USNM 123324.

Uncinocythere zancla Hobbs & Walton, 1963a. Holotype USNM 108033.

Uncinocythere zaruri Hobbs, 1971b. Holo- type USNM 128827.

Veticambarus Hobbs, 1969c.

Villalobosus Hobbs, 1972a.

Virilastacus Hobbs, 1991f.

Waltoncythere Hobbs & Peters, 1979a.

1975c.

& Walton,

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Zilchia falcata Rodriguez & Hobbs, 1989d. Holotype MNHN B20118.

Acknowledgments

Many of Hobbs’s colleagues, friends, and family contributed to this biography, and we are indebted to all of them for taking time to read a late draft and their interest, support, corrections, suggestions, and pho- tographs, all of which materially improved our effort. We acknowledge with thanks the following individuals: Fenner A. Chace, Jr., National Museum of Natural History, Smithsonian Institution, Washington, D.C.; John E. Cooper, North Carolina State Mu- seum of Natural Sciences, Raleigh; Joseph FE Fitzpatrick, Jr., University of South Ala- bama, Mobile; Richard Franz, Florida State Museum, University of Florida, Gaines- ville; Horton H. Hobbs, III, Wittenberg University, Springfield, Ohio; Brian Ken- sley and Elizabeth Nelson, National Muse- um of Natural History, Smithsonian Insti- tution, Washington, D.C.; James E Payne, Memphis State University, Tennessee; Dan- iel J. Peters, New Horizons Governor’s School for Science and Technology, Hamp- ton, Virginia; Rudolph Prins, Western Ken- tucky University, Bowling Green; Jean Pugh, Yorktown, Virginia; and Nina Hobbs Singleton, Philadelphia, Pennsylvania.

Literature Cited

All papers by Hobbs are cited above in his bibliography. They are not repeated here if cited in the text.

Chace, Fenner A., Jr. 1951. The number of species of decapod and stomatopod Crustacea.—Journal of the Washington Academy of Sciences 41(11):370-372.

Cooper, J. E., & M. R. Cooper. 1997. A new species of troglobitic crayfish of the genus Cambarus, subgenus Aviticambarus (Decapoda: Cambari- dae), endemic to White Spring Cave, Ala- bama.—Proceedings of the Biological Society of Washington 110(4):608-616.

Davidson, A. 1979. North Atlantic Seafood. New York, The Viking Press, 512 pp.

Dobbin, C. N. 1941. Fresh-water Ostracoda from Washington and other western localities.—Uni-

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versity of Washington Publications in Biology 4:174-246, 14 pls.

Fitzpatrick, J. E, Jr. 1995. Horton H. Hobs, Jr. (29 March 1914-22 March 1994).—Journal of Crustacean Biology 15(4):797—799.

. 1996. Horton H. Hobbs [sic], Jr. (29 March

1914-22 March 1994).—Crustaceana 69(2):

251-264.

. 1995. Tribute to Horton H. Hobbs, Jr (29 March 1914—22 March 1994). Pp. ix, x in M. C. Geddes, D. R. Fielder, & A. M. M. Richard- son, eds., Freshwater crayfish X. Proceedings of the International Association of Astacology Tenth Symposium 1994, Adelaide, Australia, 637 pp.

Hart, C. W., Jr. 1960. A new ostracod of the genus Entocythere from Pennsylvania.—Notulae Na- turae, Academy of Natural Sciences of Phila- delphia 335:4 pp.

503

Hoff, C. C. 1943. Two new ostracods of the genus Entocythere and records of previously described species.—Journal of the Washington Academy of Sciences 33(9):276—286.

Hoffman, R. L. 1994. +Horton Holcombe Hobbs II (1914—1994).—Banisteria 4:36—-38.

Klie, W. 1931. Crustacés Ostracodes. Campagne Speo- logique de C. Bolivar et R. Jeannel dans l’Amérique du Nord (1928), 3.—Biospeleolo- gica: Archives de Zoologie Expérimentale et Générale 71(3):333-—344.

Rioja, E. 1943. Polimorfismo femenino en los ostra- codos del género Entocythere.—Anales del In- stituto de Biologia, Universidad Autonoma de México 14(2):567—585.

Tressler, W. L. 1947. A check list of the known species of North American freshwater ostracods.—The American Midland Naturalist 38:698—707.

Wolf, B. 1934-1938. Animalium Cavernarum Catalo- gus. 2 volumes, s’Gravenhage: W. Junk. 918 pp.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

111(3):504—510. 1998.

Franconictis (Mammalia: Carnivora) from the Late Oligocene of eastern Kazakstan

Spencer G. Lucas, Robert J. Emry, and Pyruza A. Tleuberdina

(SGL) New Mexico Museum of Natural History and Science, 1801 Mountain Road N.W., Albuquerque, New Mexico 87104, U.S.A.; (RJE) Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (PAT) Institute of

Zoology, National Academy of Science, Akademgorodok, Almaty 480032, Kazakstan

Abstract.—The right dentary of a small carnivore from near Ayaguz in east- ern Kazakstan is identified as Franconictis sp. aff. F. vireti (Dehm). Francon- ictis has previously been reported only from the early Miocene of Europe. Previous reports of Plesictis from Asia are not well founded. Franconictis from Ayaguz extends the geographic range of the genus into Asia and its temporal

range back to the late Oligocene.

Fossil mammals from eastern Kazakstan are best known from the richly fossiliferous Paleogene-Neogene strata of the Zaysan ba- sin (Fig. 1). Some Paleogene and more ex- tensive Neogene mammalian assemblages are also known from the intermontane ba- sins of the northern Tien Shan drained by the Ily River and its tributaries (Tleuberdina et al. 1993, Lucas & Bayshashov 1996, Emry et al. 1997). However, between the Ily and Zaysan basins, relatively few Ce- nozoic fossil mammals are known, in part because outcrops are limited in the relative- ly low topography of the Balkash-Alakol drainage basin. Here, we add to this sparse record a jaw of the mustelid carnivore Franconictis, collected near Ayaguz. Fran- conictis has previously been known only from the early Miocene of Europe (Wolsan 1993, Morlo 1996); this specimen is the first record of the genus from Asia and ex- tends its temporal range back to the late Oligocene.

Abbreviations used.—When used in den- tal notations, upper case letters denote up- per (skull) teeth and lower case letters de- note lower (dentary) teeth. Institutional ab- breviations used are: AMNH—American Museum of Natural History, New York;

KAN-Kazak Academy of Sciences, Alma- ty.

Systematic Paleontology

Order Carnivora Bowdich, 1821 Family Mustelidae Fischer von Waldheim, 1817 Genus Franconictis Wolsan, 1993 Franconictis sp. aff. F. vireti (Dehm 1950)

Fig. 2

Referred specimen.—KAN 401-67, hor- izontal ramus of right dentary with pl—ml, from lower Miocene strata termed “Aral svita’’ near Ayaguz, Kazakstan.

Description.—The dentary is narrow and shallow with two mental foramina on its la- bial aspect—one under the anterior root of the p2 and the other under the center of the p3. The ascending ramus is broken, but ap- pears to have been tall and thin, with the anterior border of the coronoid fossa sharp- ly defined.

There is a dorso-ventrally oval alveolus for the canine at the anterior edge of the dentary as preserved. Evidently the postca- nine diastema was either short or nonexis- tent. The pl is a small, well worn trenchant tooth with a single root. The crown consists

VOLUME 111, NUMBER 3

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Fig. 1. Ily basin in the eastern part of the country.

of an anteriorly situated main cusp (proto- conid) with a long posterior surface sloping to a low heel. The p! length = 2.4 mm, width = 1.3 mm.

The p2 has two distinct roots and a trenchant crown dominated by the tall pro- toconid. There is a very small anterior cin- gulid and a low, broad heel defined by the lingual cingulid which continues around the posterior margin of the tooth. The p2 length = 3.3 mm, width = 1.6 mm.

The p3 is similar to the p2 but larger and

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has a relatively longer anterior slope. A very slight swelling low on the posterior slope of the protoconid is an incipient post- protoconid cuspid. The p4 has a similar overall shape, except the anterior cingulid bears a distinct cuspid, there is a small cus- pid (?metaconid) on the posterior slope of the protoconid, and the posterolingual cin- gulid is a sharp ridge produced into a small posterior accessory cuspid. The p3 length = 3.8 mm, width = 1.8 mm; p4 length = 4.6 mm, width = 2.1 mm.

506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. m1, occlusal (A), lingual (B) and labial (C) views. Bar scale = 5 mm.

The m1 is the only molar tooth remain- ing in the jaw, and it is substantially larger than the other teeth. The large, postero-la- bially canted protoconid dominates the tri- gonid. A relatively small and low metaco- nid is directly lingual to the protoconid, and a larger but lower paraconid projects an- tero-lingually, so that the trigonid basin is broadly open lingually. A carnassial notch interrupts the paracristid. The low talonid consists of a basin surrounded by a rela- tively sharp rim, which is much lower lin- gually than labially. The hypoconid is rel- atively large and the entoconid is present but poorly differentiated as a small cuspid on the posterolingual rim of the talonid; be- tween the hypoconid and entoconid are two small, bead-like cuspids. The m1 length = 6.4 mm, width = 2.9 mm.

Franconictis sp. aff. F. vireti (Dehm) from eastern Kazakstan, KAN 401-67, right dentary with pl—

The m2 is missing, but its position is in- dicated by two alveoli, the posterior of which is well up on the slope of the as- cending ramus. It was obviously a much smaller tooth than the ml.

Identification.—The presence of a single- rooted pl supports assignment of KAN 401-67 to the Mustelidae sensu Wolsan (1993). Among mustelids, its closest simi- larity is to “‘Plesictis,” particularly the rel- atively small species “‘P.”’ vireti from Win- tershof-West, Germany (e.g., Dehm 1950, figs. 92-99). Points of close resemblance include the shallow dentary, single-rooted pl, lack of a posterior cuspid on p2 or p3, m1 trigonid less than three times as long as the talonid, m1 protoconid relatively low, broad and posteriorly inclined, ml meta- conid higher than paraconid, m1 entoconid

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and entoconulid poorly differentiated, m2 two-rooted, and m2 alveoli on the ascend- ing ramus.

In a re-evaluation of the phylogeny and classification of European mustelids, Wol- san (1993) recently removed Plesictis vireti from Plesictis and placed it in the new ge- nus Franconictis. Given its strong similar- ity to P. vireti, we identify KAN 401-67 as Franconictis, but do not attempt a definite species-level identification of so incomplete a specimen. Therefore, we identify KAN 401-67 as Franconictis sp. aff. F. vireti. M. Wolsan (in litt., 1997) notes that KAN 401- 67 is slightly smaller and “‘more primitive”’ than F. vireti and probably represents a new species of Franconictis, but we believe the specimen is inadequate material upon which to base a new species.

Plesictis in Asia

To our knowledge (also see Russell & Zhai 1987, Werdelin 1996), Plesictis has been reported only twice from Asia. Ga- buniya (1964) first reported Plesictis based on an isolated canine, fragment of a distal humerus and partial metacarpal (Gabuniya 1964, figs. 17-19) from the Benara locality (late Oligocene) in western Georgia. These fossils represent a small carnivore, but are not sufficient to identify Plesictis. There- fore, we identify them as Carnivora, inde- terminate.

Mellett (1968) reported cf. Plesictis sp. from the upper part of the Hsanda Gol For- mation (Mongolia, late Oligocene) based on AMNH 21654, a right dentary with m1—2 (Fig. 3). In this specimen, m1 is much larg- er than m2 and has a long, tall trigonid with the basin broadly open lingually. Its proto- conid is large, tall and inclined postero-la- bially, and the metaconid is a much lower, pointed cuspid lingual to the protoconid. The paraconid is intermediate in size be- tween the protoconid and metaconid and is massive and projects anteriorly. A carnas- sial notch interrupts the paracristid. The tal- onid is very low and cup-like with a distinct

507

Fig. 3. AMNH 21654, Stenoplesictis constans from the Hsanda Gol Formation of Mongolia, right dentary fragment with m1—2, lingual (A) and occlusal (B) views. Bar scale = 5 mm.

hypoconid. The m1 length = 8.4 mm, width = 4.0 mm.

The m2 is a small, oval tooth with a three-cusped trigonid and a low talonid with a shelf-like posterior rim. The m2 length = 3.6 mm, width = 2.8 mm.

This specimen conforms well to Steno- plesictis, particularly in its relatively tall m1 trigonid, prevallid shear on ml, small m2 with a well developed trigonid and narrow, trenchant talonid, and lack of m3 (Dashzev- eg 1996). Indeed, AMNH 21654 closely re- sembles Stenoplesictis constans from the Hsanda Gol Formation (Matthew & Grang- er 1924, fig. 6F), to which we assign it. Therefore, there are no well founded reports of Plesictis in Asia.

Biostratigraphy and Biochronology

The locality from which KAN 401-67 was collected is on the right bank of the Ayaguz River, 29 km downstream from the town of Ayaguz, in eastern Kazakstan. The vertebrate-producing strata (Fig. 4) here rest with erosional unconformity on red beds

508 allUViUI ie cement ae Khorgos 2.22.02 2°%,0202%,5 svita Pavlodar legend svita — conglomerate ee sandstone ~—SS [] mudstone So Sa Se [__] limestone a) ‘ = metamorphic —— aS Aral Pots svita ae | Avagtz 6:00:60. ¢ gam) <= fossil pee locality ae Seeee = a Paleogene ->—————4] — 1 on u S woes we ! SVAl PaleoZzoicy | =e r= ire

Fig. 4. Measured section of strata exposed in the Ayaguz River basin showing the stratigraphic position of the Ayaguz fossil mammal locality (after Tolochko & Aubekerova 1971).

that elsewhere in the Balkash-Alakol basin produce middle Eocene (Irdinmanhan) mammals (on the Shinzhaly River to the south: Didenko-Kislitsina 1990; Lucas et al. 1997). The fossiliferous bed at the Ayaguz River locality is composed of rusty green sandy clays and sandy gravel lenses at the base of a section referred to as ‘“‘Aral svita”’ by Tolochko & Aubekerova (1971). In ad- dition to Franconictis, the following taxa have been reported from this site: the lag- omorphs Sinolagomys cf. S. major Bohlin

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and Desmatolagus cf. D. robustus Matthew & Granger; the castorids Asiacastor ante- cedens Lychev and Steneofiber depereti Mayet; and an indeterminate tragulid artio- dactyl (Musakulova 1971, Erbayeva 1982, Lychev 1982).

Desmatolagus ranges in age from Ergi- lian to Shandgolian in China-Mongolia, and is a characteristic Oligocene genus (Russell & Zhai 1987, Qiu & Qiu 1995). The och- otonid Sinolagomys is a characteristic late Oligocene taxon in China and Mongolia, though it has its youngest occurrence (S. pachygnathus) in the early Shanwangian (early Miocene) of China (Qiu & Qiu 1995). Therefore, Erbayeva (1982) assigned the Ayaguz mammal locality a late Oligo- cene age.

Lychev (1982, 1987, in Lychev & Au- bekerova 1971) described Asiacastor from localities in eastern Kazakstan that he as- signed a Miocene age, although there is no clear basis for the age assignment. Indeed, Asiacastor is known from the late Oligo- cene Aral local fauna, north of the Aral Sea in western Kazakstan (Lucas et al. 1998). In Asia, the genus Steneofiber is of Oligo- cene and Miocene age (Russell & Zhai 1987). Tragulids range through the Oligo- Miocene boundary, so the indeterminate tragulid from the Ayaguz locality is of no precise biochronological significance. We thus support Erbayeva’s (1982) assignment of a late Oligocene age to the Ayaguz mam- mal locality.

Franconictis is known in European early Miocene (MN 1—4) mammal assemblages (Mein 1989, Bruijn et al. 1992, Wolsan 1993, Morlo 1996, Steininger et al. 1996). Its occurrence in eastern Kazakstan thus is an extension of its temporal range back to the late Oligocene. Wolsan’s (in litt. 1997) interpretation of the Kazak specimen of Franconictis as more primitive than Euro- pean Franconictis vireti is consistent with an older age for the Kazak occurrence. Fur- thermore, given that the distribution of Franconictis in Asia is largely unknown, an older age for the genus in Asia than in Eu-

VOLUME 111, NUMBER 3

rope is quite plausible. On face value it might suggest an Asian origin of the genus during the Oligocene, and subsequent im- migration to Europe in the early Miocene.

Acknowledgments

The National Geographic Society (Grant 5412-95) and the Smithsonian Institution’s Charles D. Walcott Fund supported this re- search. R. Tedford, X. Wang, and M. Wol- san provided helpful comments on carni- vore taxonomy and the identification of the specimen reported here. The manuscript was improved by the careful critical re- views of J. Flynn and M. Wolsan.

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Bruijn, H. De, R. Daams, G. Daxner-H6ck, V. Fahl- busch, L. Ginsburg, P. Mein, J. Morales, E. Heinzemann, D. E Mayhew, A. J. Van Der Muelen, N. Schmidt-Kittler, & M. Telles An- tunes. 1992. Report of the RCMNS working group on fossil mammals, Reisensburg 1990.— Newsletters on Stratigraphy 26:65-118.

Dashzeveg, D. 1996. Some carnivorous mammals from the Paleogene of the eastern Gobi Desert, Mongolia, and the application of Oligocene car- nivores to stratigraphic correlation.—American Museum Novitates 3179:1—14. .

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Didenko-Kislitsina, L. K. 1990. Geologicheskoye raz- vitiye yugo-vostochnovo Kazakhstana v Kai- nozoye [Geological development of southeast- em Kazakstan during the Cenozoic]. Akademi- ya Nauk Kazakhskoy SSR, Institut Gidrogeo- logii 1 Gidrofiziki, Almaty, 49 pp.

Emry, R. J., S. G. Lucas, & B. U. Bayshashov. 1997. Brontothere bone bed in the Eocene of eastern Kazakstan.—Journal of Vertebrate Paleontology 17 (supplement to no. 3):44A.

Erbayeva, M. A. 1982. Kainozoiskie zaitseobraznye Kazakstana [Cenozoic lagomorphs of Kazak- stan].—Materialy po Istorii Fauny i Flory Ka- zakhstana 8:25-—38.

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Gabuniya, L. K. 1964. Benarskaya fauna oligotseno- vykh pozvonochnykh [Benara fauna of Oligo- cene vertebrates]. Izdatelstvo ‘‘Metsniereba,”’ Tbilisi, 266 pp.

Lucas, S. G., & B. U. Bayshashov. 1996. The giant rhinoceros Paraceratherium from the late Oli- gocene at Aktau Mountain, southeastern Ka- zakhstan, and its biochronological signifi- cance.—Neues Jahrbuch fiir Geologie und Pa- laontologie, Monatshefte 1996:539—548.

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cene Perissodactyla from the Shinzhaly River,

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, E. G. Kordikova, & R. J. Emry. 1998. Oli- gocene stratigraphy, sequence stratigraphy and mammalian biochronology north of the Aral Sea, western Kazakhstan.—Bulletin of the Car- negie Museum of Natural History 34:313-348.

Lychev, G. E 1982. Novyye nakhodki bobrovikh v Pavlodarskoi i Semipalatinskoi oblastyakh [New finds of beavers in the Pavlodar and Sem- ipalatinsk Districts].—Materialy po Istorii Fau- ny i Flory Kazakhstana 8:39—49.

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Matthew, W. D., & W. Granger. 1924. New Carnivora from the Tertiary of Mongolia.—American Mu- seum Novitates 104:1—9.

Mein, P. 1989. Updating of MN zones. Pp. 73—90 in E. H. Lindsay, V. Fahlbusch & P. Mein, eds., European Neogene mammal chronology. Ple- num Press, New York.

Mellett, J. S. 1968. The Oligocene Hsanda Gol For- mation, Mongolia: A revised faunal list— American Museum Novitates 2318:1—16.

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Musakulova, L. T. 1971. Myestonakhozhdeniye isko- payemykh tragulid v Kazakhstanye [Localities of fossil tragulids in Kazakhstan].—Materialy po Istorii Fauny i Flory Kazakhstana 5:52—56.

Qiu, Z., & Z. Qiu. 1995. Chronological sequence and subdivision of Chinese Neogene mammalian faunas.—Palaeoecology, Palaeogeography, Pa- laeoclimatology 116:41—70.

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Steininger, EF FE, W. A. Berggren, D. V. Kent, R. L. Bernor, S. Sen, & J. Agusti. 1996. Circum-Med- iterranean Neogene (Miocene and Pliocene) marine-continental chronologic correlations of European mammal units. Pp. 7—46 in R. L. Ber- nor, V. Fahlbusch, & H-W. Mittmann, eds., The evolution of western Eurasian Neogene mam- mal faunas. Columbia University Press, New York.

Tleuberdina, P A., L. T. Abdrakhmanova, & B. U. Bayshashov. 1993. Rannemiotsyenovaya fauna mlyekopitayushikh Dzhungarskovo Alatau (G. Aktau) [Early Miocene mammalian fauna of the Dzhungarian Alatau (Aktau Mountain)].—Ma-

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terialy po Istorii Fauny 1 Flory Kazakhstana 12: 92-115.

Tolochko, V. V., & P A. Aubekerova, P. A. 1971. Eot- sen-Pliotsenovyye otlozheniya sredney chasti basseyna r. Ayaguz [Eocene-Pliocene outcrops of the middle part of the Ayaguz River ba- sin].—Geologiya i Poleznye Iskopayemyye Yuzhnovo Kazakstana, 4:154—160.

Werdelin, L. 1996. Carnivores, exclusive of Hyaeni- dae, from the later Miocene of Europe and western Asia. Pp. 271—289 in R. L. Bernor, V. Fahlbusch & H-W. Mittmann, eds., The evolu- tion of western Eurasian Neogene mammal fau- nas. Columbia University Press, New York.

Wolsan, M. 1993. Phylogeny and classification of early European Mustelida (Mammalia: Carnivora).— Acta Theriologica 38:345—384.

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111(3):511—520. 1998.

Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 6. An intergeneric hybrid, Aglaiocercus kingi X Metallura tyrianthina, from Venezuela

Gary R. Graves

Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.

Abstract.—An intergeneric hybrid hummingbird, Aglaiocercus kingi X Me- tallura tyrianthina, is described. External measurements of the hybrid are in- termediate of those of the parental species. Back plumage iridescence is bluer (511 nm) in the hybrid than in either of the parental species (553-571 nm). This color shift is thought to be caused by a developmental aberrancy or mu- tation which affects melanin granules that produce iridescence in feather ker-

atins.

Under certain circumstances, interspecif- ic hybridization may be an important source of genetic exchange among avian lineages that may create favorable conditions for rapid and significant evolutionary change (Grant & Grant 1992). From an analysis of data in Panov’s (1989) catalog of avian hy- brids, Grant & Grant (1992) reported that 19.1% (61 of 319) of hummingbird species has hybridized in nature. A surprising 69.2% (36 of 52) of the hybridizing pairs is intergeneric (taxonomy of Sibley & Mon- roe 1990), a finding consistent with Prager & Wilson’s (1975) thesis that interspecific hybridization potential is slowly lost during avian evolution. The true extent of hybrid- ization among hummingbirds, however, is imperfectly known. Panov’s (1989) compi- lation includes numerous poorly document- ed or erroneous records, as did its antece- dent (Gray 1958). Moreover, many new hy- brid combinations have been reported re- cently (e.g., Graves 1990, 1996a, 1998a; Graves & Zusi 1990; Hinkelmann 1996; Weller & Schuchmann 1997). A definitive analysis of hybridization and phyletic retic- ulation must await a robust phylogeny and a systematic survey of purported hybrids, type specimens, and museum collections.

Here I describe an intergeneric hybrid com- bination, Aglaiocercus kingi X Metallura tyrianthina.

Materials and Methods

The unsexed specimen (American Mu- seum of Natural History [AMNH] 146645) was collected by S. Gabaldon in Estado Mérida, Venezuela. The exact locality, ele- vation, and date of collection are unknown. The specimen appears to be a male in sub- definitive plumage as evidenced by the faint striations on the maxillary ramphotheca (see Ortiz-Crespo 1972) and by its elongat- ed tail (Fig. 1 & 2). Five different identifi- cations have been written in ink and pencil (in quotations below) on the two attached AMNH labels since the specimen was cat- aloged in 1927 (in probable order of occur- rence): (a) ““Cyanolesbia’’ [=Aglaiocer- cus]; (b) ‘‘Aglaiocercus ?caudata”’ [=Aglaiocercus kingi caudatus]; (c) ““Me- tallura purpureicauda”’ [=Chalcostigma purpureicauda]; (d) ““?Hybrid?, Aglaiocer- cus caudata X Ramphomicron”’ [=Aglaio- cercus kingi caudatus X Ramphomicron microrhynchum]; and (e) ““Aglaiocercus emmae_ caudata, (melanistic aberration),

512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Ventral views of male Aglaiocercus kingi caudatus (top), Metallura tyrianthina oreopola (bottom), and a probable hybrid, A. kingi caudatus X M. tyrianthina oreopola (AMNH 146645).

Fig. 2. Probable hybrid, Aglaiocercus kingi caudatus X Metallura tyrianthina oreopola (AMNH 146645).

VOLUME 111, NUMBER 3

fide J. Berlioz, Apr. 1975” [=Aglaiocercus kingi caudatus}.

I compared the oft-identified specimen with series of all species in the subfamily Trochilinae, the typical hummingbirds (Zusi & Bentz 1982, Sibley & Monroe 1990, Bleiweiss et al. 1997), in the collections of the American Museum of Natural History and the National Museum of Natural His- tory (USNM), Smithsonian Institution. For the purpose of hybrid diagnosis, I consid- ered all hummingbirds (Trochilinae; taxon- omy of Sibley & Monroe 1990) that occur in Estado Mérida as potential parental spe- cies (Phelps & Phelps 1958, Meyer de Schauensee & Phelps 1978) (Appendix 1). In addition, I compared the specimens di- rectly with the holotypes of Chalcostigma purpureicauda (AMNH 483931), Lesbia ortoni (AMNH 156651), Zodalia thaumas- ta (USNM 173911), and Aeronympha pro- santis (Field Museum of Natural History, FMNH 11852), and with notes, photo- graphs, and videotape of the holotype of Heliangelus zusii (Academy of Natural Sci- ences of Philadelphia, ANSP 159261).

Color descriptions given in Appendix 2 were made under natural light. I evaluated the color of dorsal plumage (center of back) and the ventral surfaces of the rectrices with a reflectance spectrophotometer equipped with a 11.0 mm aperture (Color Mate Col- orimeter, Milton Roy). The colorimetric characters were described in terms of op- ponent-color coordinates (L, a, b) (Hunter & Harold 1987). This system is based on the hypothesis that signals from the cone receptors in the human eye are coded by the brain as light-dark (L), red-green (a), and yellow-blue (b). The rationale is that a color cannot be red and green or yellow and blue at the same time. Therefore “‘redness’’ and ““greenness”’ can be expressed as a single value a, which is positive if the color is red and negative if the color is green. Likewise, “‘yellowness”’ or “‘blueness”’ is expressed by b for yellows and —b for blues. The third coordinate L, ranging from 0 to 100, describes the “‘lightness”’ of color; low val-

513

ues are dark, high values are light. In other words, the more light reflected from the plumage the higher the L value will be. It should be noted that visual systems in hum- mingbirds (e.g., Goldsmith & Goldsmith 1979) differ significantly from those of hu- mans. The relevance of opponent color co- ordinates to colors perceived by humming- birds is unknown. Dominant wavelengths reflected from plumage surfaces are listed for comparison. Data in Table 1 were com- piled from the averages of five independent measurements (specimen moved from ap- erture between trials) for each plumage area per specimen.

Measurements of wing chord, bill length (from anterior extension of feathers), and rectrix length (from point of insertion of the central rectrices to the tip of each rectrix) were taken with digital calipers and round- ed to the nearest 0.1 mm (Table 2). Mea- surements and least squares regression lines were projected on bivariate plots to illus- trate size differences (Wilkinson 1989).

There are three alternatives to consider— the specimen represents an aberrant color morph of A. kingi or some other species, a hybrid, or an undescribed species. The specimen differs significantly in size and shape from all species in Appendix 1. In particular, the rectrices of the specimen are considerably wider, flatter in cross section, and more iridescent on the ventral surfaces than in A. kingi, indicating that it is not sim- ply a melanistic example of that species as suggested by Berlioz on the specimen label. Because hybrids have no standing in zoo- logical nomenclature, the burden of proof rests on the investigator to refute this pos- sibility before bestowing species status on a unique specimen. Because the evidence points to hybridization, I refer to the spec- imen as a hybrid in the remainder of the paper.

The diagnosis was approached in a hi- erarchical manner. The presumed parental species of the hybrid first were hypothe- sized through the comparative analysis of plumage pattern, as well as from feather

514 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Ranges and means (+standard deviation) of opponent color coordinates (L, a, b) and dominant wavelength reflected from dorsal plumage (center of back) and the ventral surface of rectrices in male Aglaio- cercus kingi caudatus, Metallura tyrianthina oreopola, and their probable hybrid (AMNH 146645).

A. kingi M. tyrianthina

Variables (n = 12) (n = 12) Hybrid

Back plumage

L (Lightness) (L) 23.6—30.1 21.5—26.6 24.1 PH 2e Wess DAD r= eA)

a (Red [+]/Green [—]) (a) —14.8-(—6.6) —7.1-(—0.6) —7.9 =10:9 = 2-9 S37 3E i183

b (Yellow [+]/Blue [—]) (b) 15.4—20.2 12.6—20.9 2.9 17.8 + 1.4 17.3 2 DB

Dominant Wavelength (nm) 553.1-563.8 562.8—570.5 511.3 Sas) 2 SD) 566.7 + 2.6

Ventral surface of rectrices

L (Lightness) (L) 18.8—21.1 20.3—23.9 20.6 2 On OM Die = NAD)

a (Red [+]/Green [—]) (a) 3.8-5.9 11.5-17.0 10.6 49 + 0.6 14.6 + 1.6

b (Yellow [+]/Blue [—]) (b) —4.3-1.5 5.5-14.9 =7/.\l —0.9 + 1.9 OS) a6 DY)

Dominant Wavelength (nm) 487.2-652.1 588.9-622.7 541.0

SAO 22 47/7) 602.5 + 11.2

and bill shape. The restrictive hypothesis then was tested with a quantitative analysis of size and external proportions. Concor- dance of results is regarded as strong sup-

Table 2.—Ranges and means (+standard deviation) of measurements (mm) of males of Aglaiocercus kingi caudatus (subdefinitive plumage, see Appendix 2), Metallura tyrianthina oreopola, and their presumed hybrid (AMNH 146645).

port for the hypothesis (Graves 1990, Graves & Zusi 1990).

Results and Discussion

Plumage characters.—Salient characters of the hybrid that permit its parental species to be identified include: (a) moderately elongated outer rectrices (fork depth = 23.7 mm), nearly flat in cross section; (b) un- marked rectrices exhibiting metallic irides-

A. kingi M. tyrianthina cence on the dorsal and ventral surfaces; (c) Character (n = 21) (n = 20) Hybrid Sig i short tibial plumes (not extending to hal- Wing chord 59.7-63.6 54.0-61.5 61.4 yx); and (d) short straight bill (11.1 mm). GiEsi== sO Sell 25 DP ies : Two species in the pool of potential paren- Bill 11.0-13.6 91-106 11-1 1 d A = OF Ao aati tal species (Appendix 1) possess elongate i i > : - Rectrix | DD BEDS 33.7389 "32, ‘alls Cengiheiitect > ae 241 +08 261 £16 tus underwoodii and Aglaiocercus kingi. Rectrix 2 WO.O-33.6 34.3-41.3 36.9 Ocreatus can be deleted from the list of Sills} se 37.9 + 1.8 possibilities because the hybrid lacks evi- Rectrix 3 38.9-46.7 36.0-43.6 43.1 dence of spatulate rectrices or lengthened 42.8 + 1.9 Sie) 25 1 tibial plumes. Aglaiocercus kingi is thus Rectrix 4 53.7-61.5 38.4-45.3 49.0 identified as one of the parental species. 20) Zot AUS = 2:1 Determination of the other parental spe- Ege 3 G75 L027 37.9-46.3 55.8 cies is equally straightforward. The inten- 3 oe) ADD, se 7)

sity of the metallic iridescence reflected

VOLUME 111, NUMBER 3

600

S4) @ oOo

560

540

520

Dominant Wavelength (nm) of Back Plumage

500

Lightness (L) of Back Plumage

Fig. 3.

515

Yellowness (b) of Back Plumage

O -10 -5 O 5 10 15

Yellowness (b) or Blueness (-b) of Ventral Surface of Rectrices

Bivariate plots of spectrophotometric data from male hummingbirds: Aglaiocercus kingi caudatus

(circles); Metallura tyrianthina oreopola (diamonds); and a probable hybrid, A. kingi caudatus * M. tyrianthina

oreopola (triangle; AMNH 146645).

from the ventral surfaces of the hybrid’s rectrices is matched or exceeded only in Metallura tyrianthina. Details of plumage pattern and feather shape are sufficient to suggest that the parentage of the hybrid is Aglaiocercus kingi X Metallura tyrianthina (see Appendix 2). None of the other species in Appendix 1, considered two at a time, can account for the characters observed in the hybrid. In particular, the ventral rectri- cial surfaces of the hybrid are metallic red- dish-purple as opposed to dull black or pur- plish-black in both Ramphomicron micro- rhynchum and A. kingi, effectively elimi- nating this pair of species from contention.

The question of plumage color.—Irides- cence in hummingbirds is caused by the in- _terference of light reflected from the upper and lower surfaces of gas-filled vacuoles in melanin granules in the keratin of feather barbules, which are compactly stacked in 7—15 layers in the barbule keratins (Dorst 1951; Greenewalt et al. 1960a, 1960b; Lu- cas & Stettenheim 1972). Carotenoid pig- ments have not been extracted from irides- cent feathers. Employing transmission electron microscopy and micro-spectropho- tometry, Greenewalt et al. (1960a, 1960b) found melanin granules to be elliptical in shape, about 2.54 long, 1.54 wide, and

0.15y thick. Briefly summarized, they found that granules contain a fairly uniform layer of gas-filled vacuoles that resemble a monolayered foam. The melanin matrix and gas-filled vacuoles have refractive indices of ~2.0 and 1.0, respectively. The color of iridescence varies according to the thick- ness of the granule and the amount of gas in the vacuoles. Iridiscent colors change from blue to green to orange and finally to red, as the effective refractive index of granules advances from 1.45 to 1.90 (figure 4 of Greenewalt et al. 1960a). Melanin granules in noniridescent parts of feathers lack vacuoles.

The pattern of bluish-green iridescence in the hybrid corresponds precisely to that of green iridescence in the parental species, suggesting a single mutational or develop- mental aberrancy that affects plumage col- or. The dominant wavelength reflected from dorsal plumage is shorter in the hybrid (511 nm) than in the parental species: Aglaio- cercus kingi (553-564 nm) and Metallura tyrianthina (563-571 nm) (Table 1, Fig. 3). The premise that “‘hybridization produces no traits characteristic of genera or species other than those involved in the particular cross”’ (Banks & Johnson 1961:3) was ex- tended to spectrophotometric measures of

516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

65

60

Wing Length

55

50 20 40 60 80

Length Rectrix 5

40

35

30

Length Rectrix 1

25

20 20 30 40

Length Rectrix 2

Length Rectrix 1

30 40 50

Length Rectrix 4

Fig. 4. Bivariate plots of mensural characters of male hummingbirds: Aglaiocercus kingi caudatus (triangles); Metallura tyrianthina oreopola (circles); and a probable hybrid, A. kingi caudatus X M. tyrianthina oreopola

100

60

120

70

14

13

12

Bill Length

11

10

20 40 60 80 100 120 Length Rectrix 5

40

w@ (o)

() Oo

Length Rectrix 1

i) a

20 35 40 45 50

Length Rectrix 3

40

ie) on

ie) Oo

Length Rectrix 1

ie) 6) a

Length Rectrix 5

(diamond; AMNH 146645). Least squares regression lines are projected for comparison.

er

VOLUME 111, NUMBER 3

plumage color (Graves 1996b). Both as- sumptions are violated in the present ex- ample.

Elucidation of the micro-structure and spectrophotometric properties of melanin granules in Aglaiocercus kingi, Metallura tyrianthina, and the hybrid is beyond the scope of this paper. Several observations on hummingbird plumages, however, are worth noting. A variety of plumage aber- rations, unassociated with hybridization, has been observed in hummingbirds, in- cluding leucism, albinism, schizochroism, erythrism, and melanism (Salvin 1892, Banks & Medina 1963, Greenway 1978, Graves 1998b). Subtle within-population variation in iridescent color is commonly observed whenever large series of species are assembled. Although post-mortem ef- fects may be partially responsible in some cases (Graves 1986, 1991), most of the ob- served variation in iridescence among in- dividuals, factoring out the effects of sex and age (see Bleiweiss 1992), is due to ge- netic and developmental factors. Pro- nounced color shifts of the magnitude ob- served in this hybrid are rare but not un- known (e.g., Salvin 1892, Greenway 1978). The example described here seems to be the first in which a hybrid hummingbird exhib- its a major plumage aberrancy.

External measurements.—One of the guiding principles of hybrid diagnosis is that hybrids are not larger or smaller than their parental species (Graves 1990). Mor- phological luxuriance or dwarfism in hybrid hummingbirds has not been recorded. Male Aglaiocercus kingi and Metallura tyrianthi- na are similar in bill length (cumulative range, 9.1—13.6 mm) and wing chord (cu- mulative range, 54.0—63.6 mm), but differ markedly in tail size and shape (Table 2, Fig. 4). Bivariate plots of rectrix length of the parental species exhibit positive (1 vs. 2) or negative (1 vs. 3, 1 vs. 4, 1 vs. 5) allometry. Except for rectrix 3, measure- ments of the hybrid fall between the char- acter means for A. kingi and M. tyrianthina, and, in several cases, approximate the val-

S17)

ues predicted by least squares regression on bivariate plots (Table 2, Fig. 4). In sum- mary, plumage pattern, distribution and in- tensity of iridescence, rectrix shape, and mensural characters provide strong support for the hypothesis of hybridity (Aglaiocer- cus kingi X Metallura tyrianthina). Previous records.—A hybrid of Aglaio- cercus kingi and Metallura tyrianthina was reported once before by Meyer de Schauen- see (1947:108), who described a specimen (No. 134) obtained in Bogota, circa 1909, from the Brother Nicéforo Maria collection:

“... fore-crown glittering brassy green, hind crown and back dark bluish green, rump and upper tail coverts bluer; chin dusky, throat patch shaped as in Metallura tyrianthina but blue instead of green; breast dark bluish green, bases and edges of the feathers buffy; belly dark green, the bases of the feathers white, showing through and giving a some- what barred appearance; tail purple, deeply forked, the outermost tail feathers 50 mm., the central ones 30 mm., wing 61 mm., culmen 12.5 mm.”

The brief description of Nicéforo’s spec- imen differs in minor details from the Ven- ezuelan specimen (AMNH 146645). The two specimens are similar in size. Nicéfo- ro’s specimen possesses a bluish gorget as might be expected in an adult male hybrid of Metallura t. tyrianthina and Aglaiocer- cus k. kingi from the Cordillera Oriental of the Colombian Andes. Whereas I charac- terized the back color of the Venezuelan specimen as “greenish-blue,’’ Meyer de Schauensee used the term “bluish-green’”’ for Nicéforo’s specimen. This and other dis- crepancies might reflect semantics or real differences in color. Unfortunately, the whereabouts of Nicéforo’s specimen is un- known, although another mentioned in Meyer de Schauensee’s paper was deposit- ed in the Academy of Natural Sciences of Philadelphia (Nicéforo no. 148, now ANSP 159261; Graves 1993).

Acknowledgments

I thank Richard Banks, Robert Bleiweiss, Kenneth C. Parkes, and Richard Zusi for critiques of the manuscript. I thank the cu-

518

rators and staffs of the American Museum of Natural History, New York, the Field Museum of Natural History, Chicago, and the Academy of Natural Sciences of Phil- adelphia, for permitting me to examine specimens in their care and for specimen loans. Photographic prints were provided by Smithsonian photographic services. Mu- seum work was supported by the Alexander Wetmore Fund and the Department of Ver- tebrate Zoology, National Museum of Nat- ural History, Smithsonian Institution.

Literature Cited

Banks, R. C., & N. K. Johnson. i961. A review of North American hybrid hummingbirds.—Con- dor 63:3—28.

, & D. R. Medina. 1963. An albinistic Anna Hummingbird.—Condor 65:69—70.

Bleiweiss, R. 1992. Reversed plumage ontogeny in a female hummingbird: implications for the evo- lution of iridescent colours and sexual dichro- matism.—Biological Journal of the Linnean So- ciety 47:183-195.

, J. A. W. Kirsch, & J. C. Matheus. 1997. DNA hybridization evidence for the principal lineages of hummingbirds (Aves: Trochilidae).—Molec- ular Biology and Evolution 14:325—343.

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Goldsmith, T. H., & K. M. Goldsmith. 1979. Discrim- ination of colors by the black-chinned hum- mingbird, Archilochus alexandri.—Journal of Comparative Physiology A 130:209—220.

Grant, P. R., & B. R. Grant. 1992. Hybridization of bird species.—Science 256:193—-197.

Graves, G. R. 1986. Systematics of the Gorgeted Woodstars (Trochilidae: Acestrura).—Proceed- ings of the Biological Society of Washington 99:218—-224.

. 1990. Systematics of the ““green-throated sun-

angels” (Aves: Trochilidae): valid taxa or hy-

brids?—Proceedings of the Biological Society

of Washington 103:6—25.

. 1991. Taxonomic status of the Sword-billed

Hummingbird (Ensifera ensifera caerules-

cens).—Bulletin of the British Ornithologists’

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. 1993. Relic of a lost world: a new species of

sunangel (Trochilidae: Heliangelus) from Bo-

gota.—Auk 110:1-8.

. 1996a. Diagnoses of hybrid hummingbirds

(Aves: Trochilidae). 2. Hybrid origin of Erioc-

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109:373-—390.

, 1998a. Diagnoses of hybrid hummingbirds

(Aves: Trochilidae). 5. Probable hybrid origin

of Amazilia distans Wetmore & Phelps.—Pro-

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, 1998b. Taxonomic notes on hummingbirds

(Aves: Trochilidae). 1. Eriocnemis dyselius El-

liot, 1872 is a melanistic specimen of Eriocnem-

is cupreoventris (Fraser, 1840).—Proceedings

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, & R. L. Zusi. 1990. An intergeneric hybrid hummingbird (Heliodoxa leadbeateri X Helian- gelus amethysticollis) from northern Colom- bia.—Condor 92:754—760.

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Greenewalt, C. H., W. Brandt, & D. D. Friel. 1960a. Iridescent colors of hummingbird feathers.— Journal of the American Optical Society 50: 1005-1016.

. 1960b. The iridescent colors of hummingbird feathers.—Proceedings of the American Philo- sophical Society 104:249—253.

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, & W. H. Phelps, Jr. 1978. A guide to the birds of Venezuela. Princeton University Press, 424 Pp.

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ical isolation in birds (in Russian). Nauka, Mos- cow, 510 pp.

Phelps, W. H., & W. H. Phelps, Jr. 1958. Lista de las aves de Venezuela con su distribuci6n. Tomo 2, Parte 1. Editorial Sucre, Caracas, 317 pp.

Prager, E. M., & A. C. Wilson. 1975. Slow evolution- ary loss of the potential for interspecific hybrid- ization in birds: a manifestation of slow regu- latory evolution.—Proceedings of the National Academy of Science USA 72:200—204.

Salvin, O. 1892. Catalogue of the birds in the British Museum, Vol. 16, London, 703 pp.

Sibley, C. G. & B. L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Uni- versity Press, New Haven, Connecticut, 1111 Pp.

Weller, A.-A., & K.-L. Schuchmann. 1997. The hybrid origin of a Venezuelan Trochilid, Amazilia dis- tans Wetmore & Phelps 1956.—Ornithologia Neotropical 8:107—112.

Wilkinson, L. 1989. SYSTAT: the system for statistics. SYSTAT, Inc., Evanston, Illinois, 822 pp. Zimmer, J. T. 1952. Studies of Peruvian birds. No. 62. The hummingbird genera Patagona, Sappho, Polyonymus, Ramphomicron, Metallura, Chal- costigma, Taphrolesbia, and Aglaiocercus.—

American Museum Novitates 1595:1—29.

Zusi, R. L., & G. D. Bentz. 1982. Variation of a muscle in hummingbirds and swifts and its systematic implications.—Proceedings of the Biological Society of Washington 95:412—420.

Appendix 1

Species of hummingbirds that occur in Estado Mé- rida, Venezuela: Campylopterus falcatus, Colibri thal- assinus, C. coruscans, Klais guimeti, Lophornis delat- trei, L. stictolophus, Chlorestes notatus, Chlorostilbon mellisugus, C. poortmani, Thalurania furcata, Hylo- charis cyanus, Chrysuronia oenone, Amazilia versi- color, A. fimbriata, A. viridigaster, Chalybura buffonii, Heliodoxa leadbeateri, Sternoclyta cyanopectus, Coe- ligena coeligena, Ocreatus underwoodii, Aglaiocercus kingi, Heliomaster longirostris, Chaetocercus jourda- nil.

Appendix 2

Comparative description of plumages of male Aglaiocercus kingi caudatus, Metallura tyrianthina or- eopola, and their presumed hybrid, AMNH 146645. The molts and plumages of male Aglaiocercus spp. are incompletely known. Young males (>6 months?) ac- quire a plumage that differs from the definitive plum- age of adult males. This subdefinitive plumage is char- acterized by shorter outer rectrices, an incompletely developed crown patch (8 of 20 examined), and rem- nants of a white rump patch (see Zimmer 1952). One quarter (5 of 20) of the males in subdefinitive plumage

519

retain a few striations on the maxillary ramphothecum, a character usually interpreted as a sign of immaturity (Ortiz-Crespo 1972). The descriptions of Aglaiocercus kingi given below refer to the subdefinitive plumage.

Descriptions of structural colors are unusually sub- jective, as color seen by the observer varies according to the angle of inspection and direction of light. For this reason I use general color descriptions.

The dark bluish-green crown of young kingi is re- placed (from anterior to posterior) by an ovate crown patch composed of brilliant bluish-green feathers. The hindneck, back, and rump are dark green; feathers are gray, tipped with green. Upper-tail coverts are bluish- green. A few white feathers form an indistinct patch on the lower back.

The dorsal plumage of tyrianthina is dark dusky green, brighter on the crown, and with coppery high- lights on the lower back and rump. Feathers are gray, banded subterminally with coppery-green, and tipped broadly with dark green. When viewed head-on in di- rect light, plumage posterior to the midcrown region appears sooty black. Immature tyrianthina lack a con- trasting rump patch.

Under a diffuse light source, the dorsum of the hy- brid is a rich greenish-blue (paler on the crown), a color that is distinctly different from that of the pre- sumed parental species. Feathers on the left side of the forecrown are discolored, possibly by a preservative chemical. Dorsal feathers are dark gray, tipped with greenish-blue. Crown feathers are not modified as in adult kingi. When light is reflected obliquely (>90 from the observer), the dorsal plumage appears purple; when viewed head-on the hindcrown, back, and rump appear black. A few rump feathers are tipped with buff.

The ventral plumage of kingi is medium green ex- hibiting subdued iridescence. A few small shining green disks occur on the throat of more mature indi- viduals. The barbs of ventral feathers are narrowly tipped with buff or grayish-buff, especially along the midline of the abdomen. Some males in juvenile and subdefinitive plumage (e.g., AMNH 484067) have a white or buffy-white line extending from the base of the bill posterior to below the eye. Downy vent feath- ers are dark gray tipped with white or pale gray. Un- dertail coverts are dark green broadly edged with buff. Tibial feathers are short (extending half way to the hallux from the tibiotarsal joint), dark olive-gray and narrowly tipped with scattered grayish-buff barbs.

The venter of tyrianthina is dark dull green; feathers are tipped with buff or grayish-brown, especially along the midline. Feather tipping imparts a mottled appear- ance to the underparts. A narrow ovate gorget extends from the chin to the upper breast in tyrianthina. When view head-on in direct light, the auriculars and sides of the throat appear matte black and contrast greatly with the brilliant green gorget. Subadult males have a buffy line extending from the bill to below the eye.

520

Vent feathers are dark gray tipped with white or pale gray. Under-tail coverts are buff with a large broadly lanceolate spot (coppery- or bronzy-red) along the midline. Tibial feathers (dark gray tipped with buff) extend to the base of the hallux.

The venter of the hybrid is similar in pattern to kin- gi, but with the green portions replaced with bluish- green (paler than dorsum). A small brilliant feather occurs along the midline at midthroat (deep blue tip separated from the gray base by narrow blending bands of light blue and coppery-gold). A buffy-white stripe extends from the bill to below the eye (Fig. 2). Vent feathers are dark gray tipped with white. Undertail co- verts are buff with a lanceolate subterminal spot (pur- ple) near the midline. Tibial feathers (dark gray broad- ly tipped with buff) of the hybrid extend about halfway to the hallux, but may have been damaged by knotting of the specimen label string.

The remiges of kingi are brownish-black faintly tint- ed with purple. The outer vanes of the primary coverts and the innermost secondaries are edged with shining green or bluish-green; secondary coverts are broadly tipped with bluish-green. The remiges of tyrianthina are very similar in color but faintly tinted with bronze or olive. Wing coverts and the innermost secondaries are bronzy-green.

The remiges of the hybrid resemble those of kingi. Wing coverts and innermost secondaries are purple tipped with dark bluish-green.

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The tail of kingi is deeply forked. The basal portions of the rectrices that are obscured in the folded tail are black. The exposed dorsal sections of the inner rectri- ces (1—4) are deep brilliant purple tipped with bluish- green. The outermost rectrices (5) lack bluish-green tips, are greatly elongated, narrow (S—6 mm wide, 25 mm from tip), and bowed in cross-section. Inner rec- trices are smoothly tapered; rectrix 5 is bluntly tipped. Ventrally, the vanes are dull purplish-black. The ra- chises in kingi are blackish-brown dorsally, medium brown ventrally.

The tail of tyrianthina is shallowly forked. Rectrices are wide (10-11 mm), nearly flat in cross-section, abruptly truncate at the tip, and metallic coppery-red above and below. Rachises are dark brown above and below.

The tail of the hybrid is moderately forked. Feather size and shape are intermediate between kingi and ty- rianthina. The outermost rectrices (5) are slightly bowed in cross-section (ca. 9.3 mm at widest point). Rectrices are metallic reddish-purple, above and be- low, the innermost (1—2) are diffusely tipped with pur- ple. This iridescence, especially from the ventral sur- faces, is similar in visual essence to that of tyrianthina. Rachises are dark brown above, medium brown ven- trally.

Bill color is black in kingi, tyrianthina, and hybrid. In dorsal profile, the bills of both parental species are abruptly tapered, more so in kingi. The bill profile of the hybrid is similar to that of tyrianthina.

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111(3):521—530. 1998.

A new species of Alsodes (Amphibia: Anura: Leptodactylidae) from southern Chile

J. Ramon Formas, César Cuevas, and José Nufiez

Instituto de Zoologia, Universidad Austral de Chile, Casilla 567, Valdivia, Chile

Abstract.—A new species of frog, Alsodes kaweshkari, is described from the temperate Nothofagus forests of South America in Southern Chile. From the karyological point of view this species is included in the monticola group (2N

= 26) of the genus Alsodes.

The genus Alsodes Bell 1843, is distrib- uted in central and southern Chile and along the eastern slope of the Andes, in Argentina (south of Mendoza city). Frost (1985) in- cluded the following species in the genus: A. barrioi, A. gargola, A. illotus, A. laevis, A. montanus, A. monticola, A. nodosus, A. pehuenche, A. tumultuosus, A. vanzolinii, and A. verrucosus. Formas (1989) studied the identity and synonymy of the Chilean frog Eupsophus vittatus and concluded that this species must be included in the genus Alsodes as A. vittatus. The taxonomic status of A. illotus is not clear (Cei 1980) and ac- cording to Frost (1985) A. laevis has not been collected since its description by Phi- lippi (1902). The most remarkable charac- teristic of the males of this genus is the presence of thorny structures on the fingers and round spiny patches on the chest.

During the course of collecting frogs in southern Chile we found a new species of Alsodes which is here described. Moreover, we describe the karyotype and C-banded chromosomic pattern of the new taxon. The bands of hemoglobin of the new species were characterized electrophoretically and compared with those of A. monticola (type species of the genus).

Methods and Materials

Specimens were deposited in the Instituto de Zoologia, Universidad Austral de Chile (IZUA). Adults were measured with a dial

caliper to the nearest 0.1 mm according to Cei (1962). The following measurements were taken: snout—vent length (SVL), head length, head width, nostril-snout distance, tibia length, and foot length. Internarial dis- tance was measured according to Cei (1980) and eye diameter according to Duellman (1970).

The chromosomal characteristics were obtained from the holotype (IZUA 1624) according to the following methods: the an- imal was treated with 0.1% colchicine for two hours, then anaesthetized with diethyl ether, cut open ventrally under sterile con- ditions and its intestine carefully removed. Metaphase plates were obtained by squash- ing intestinal epithelium fragments that were hypotonically treated with distilled water, then fixed in acetic-alcohol (1:3), and finally placed in 45% acetic acid. Small tis- sue fragments were squashed between a glass slide and cover slip and dipped in liq- uid nitrogen; thereafter, the cover slip was removed with a razor blade to allow chro- mosomes to air dry. After 3 days, the chro- mosomes were stained for 15 minutes in Sorensen’s phosphate buffer (pH 6.8), con- taining 4% Giemsa solution (Formas 1991). Centromeric positions were determined ac- cording to Levan et al. (1964). Secondary constrictions were included in the measure- ments. Chromosomes were stained to reveal C-band pattern position (Sumner 1972). Microscopic slides and the specimen (IZUA

S22

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Fig. 1. Alsodes kaweshkari Holotype, IZUA 1624.

1624) were deposited in the amphibian col- lection of the Instituto de Zoologia, Univ- ersidad Austral de Chile.

Hemoglobin was obtained according to the following methodology: blood was col- lected by cardiac puncture from the holo- type and a specimen of Alsodes monticola (control species) using a sterile syringe with 3.8% of sodium citrate as anticoagulant; the red cells were washed 3 times with 0.85% NaCl and centrifuged at 700 g for 5 minutes at room temperature (20°C). The packed cells were hypotonically treated with dis- tilled water (10 volumes) for 30 minutes and centrifuged at 3500 g for 10 minutes. The supernatant was stored at —70°C. The hemolyzate was treated according to the method described by Muir (1981) with modifications. We used a minicolumn (5 X 70 mm) of Sephadex G-50 equilibrated with 1.0 mM of potassium phosphate (pH 7.0). Fractions of 500 wl were collected and their absorbances measured at 410 nm, the wave length at which the porphyrin ring of the hemoglobin molecule absorbs specifi- cally. Those fractions with the highest ab- sorbance value were used for the electro- phoretic analysis of the hemoglobin of the

M. Nowaste . 97

2cm

new species and A. monticola. To proceed with the electrophoresis the samples were denatured by heating (100°C, 5 minutes) in presence of 2% of betamercaptoethanol. Electrophoresis of the hemoglobins from Alsodes kaweshkari and A. monticola was carried out individually in vertical slab gels of polycrilamide (15%) in denaturant con- ditions at 20 mA for 20 hours (Laemmeli 1970). Gels were stained with Coomassie Blue R-250 for 2 hours and destained with acetic acid (7%). The molecular weight of the hemoglobin bands was estimated using a protein standard for molecular weight (Sigma Co).

Systematics

Alsodes kaweshkari, new species. Fig. 1

Holotype.—IZUA 1624, an adult male collected by J. Ram6n Formas on 8 March 1995, at Puerto Edén (49°8'S, 74°25'W, 10 m), Wellington Island, Ultima Esperanza Province, Magallanes and Chilean Antartic Region (XII Regién), Chile (Fig. 2).

Paratype.-—IZUA 1625, an adult male collected by Pablo Corti on 21 Oct 1995,

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46°

Peninsula de

Taitao Sf

B

(e) oO u O > ¢ Q

OCEANO

Puerto

Isla a

Wellington ~—-”

Fig. 2. Type locality of Alsodes kaweshkari.

at Seno Huemules (48°43’'5S, 74°25’W, 8 m), Ultima Esperanza Province, Magallanes and Chilean Antartic Region (XII Region), Chile (Fig. 2).

Diagnosis.—Alsodes kaweshkari can be distinguished from its congeneric species by the following combination of characters: SVL 56.5—62.2 mm; toes totally fringed; webbing of feet present between all toes, but reduced; dorso-lateral surfaces granular; skin around vent and posterior thighs gran- ular.

Description.—Based on the type series.

523

) REPUBLICA

? ARGENTINA

Seno Huemules

Body robust, arms and legs well developed. Head depressed, slightly wider than long, its length 29% of snout—vent length. Snout truncated from above, rounded in lateral profile; canthus rostralis slightly rounded, loreal region concave in cross section; nos- trils anterolateral, midway between tip of snout and anterior border of eye; eye di- ameter greater than distance between eye and nostril; internarial distance less than in- terorbital distance. Tympanum absent; post- ocular fold evident, reaching insertion of arm. Tongue rounded, without notch at tip.

524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

SSP rad ey j iis

Fig. 3. Morphological details of the holotype. (A) Palmar view of the right hand. (B) Plantar view of the

right foot.

Choanae oval-shaped; dentigerous process- es of vomer between choanae. Forelimbs of males robust. Fingers in order of increasing length: II, I, IV, II. Webbing of hand ab- sent. Fingers long with moderately globular

tips. Inner palmar tubercle ovoid, outer pal- mar tubercle rounded; subarticular tubercles rounded, (Fig. 3A). First finger with strong thorny excrescences; second finger with narrow band of spines (Fig. 4B). Palmar

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S25)

5mm

Fig. 4. Secondary sexual characters of the holotype. (A) Ventral view of the chest and throat. (B) Dorsal

view of the left hand.

surface with few thorns irregularly distrib- uted. Spines also present at lower border of mandible (Fig. 4A). Toes long, thick and fringed; in order of increasing length: I-II- III-V-IV; tips moderately rounded. Webbing present, thick, but reduced. Inner metatarsal tubercle oval, elongate; outer tubercle smaller than the inner, oval. Tarsal fold present; reaching middle of tarsus. Flanks and dorsal surface granular; ventral skin smooth with minute granules; skin around vent and posterior thighs granular. Chest of males with two bilateral rounded black patches of keratinous spines (Fig. 4A). Dor-

Table 1.—Measurements (mm) of the type series of Alsodes kaweshkari.

Holotype Paratype IZUA 1624 IZUA 1625

Character male male Snout—vent length 62.2 59.9 Head length 17.7 16.4 Head width 24.2 22.8 Eye diameter 7.0 6.7 Nostril—snout distance 4.0 3.6 Internarial distance So 7/ 5.4 Tibia length MA 27.4 Foot length 33.8 30.9

sal surfaces of head and areas below dor- solateral fold with granules.

The measurements of the holotype and the paratype are given in Table 1.

Coloration.—In alcohol, dorsal surfaces of ground, arms and legs dark gray. Venter whitish, gular area dark gray. Arms and legs whitish ventrally. The granular surface around posterior thighs dark gray. In life, dorsal surface, arms and legs brown. Belly whitish yellow. Areas around nostrils, lips and eyes light brown. The iris with a cop- pery reticulum.

Distribution and ecology.—The species is known from two Chilean localities (Puer- to Edén and Seno Huemules) (Fig. 2). The type locality (Puerto Edén) is a small beach of the Wellington Island, near a small fish- ing town. There are small forests (Notho- fagus betuloides, Embothrium coccineum, Maytenus magellanicus, and Drymis win- teri) surrounded by typical tundra with plants of the genera Donatia, Astelia, and Azorella and rushes of the families Cyper- aceae and Juncae. This area is situated in the oceanic cold temperate region (di Castri 1968). The annual mean temperature of this region is 8.8°C; the relative humidity is

526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

ey i a

ax BA aust 1

eA if 11 12

Mon

cs 8 a 8

9 10 Bk aks} 10 um } 5 $ = & a1 . 3 4 5

* A 13

Fig. 5. Chromosomes of the holotype. (A) Karyotype. (B) C-banded karyotype.

87%, and the rain fall ranges from 2000— 2500 mm per annum. The frog was col- lected under a log between the forest and the tundra. The male holotype shows its re- markable thorny excrescences on the chest

and fingers (Fig. 4A, B). The following am- phibians were also collected in this area: Bufo variegatus, Batrachyla antartandica and Eupsophus calcaratus. Lynch (1975) pointed out the presence of Atelognathus

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Table 2.—Relative length, arm ratio (mean and stan- dard deviation), and type of chromosomes (m = meta- centric; sm = submetacentric; st = subtelocentric) of metaphase chromosomes of Alsodes kaweshkari. Rel- ative length was calculated according to Bogart (1970). Arm ratio was calculated by dividing the short arm into the long arm. *Chromosomes with secondary con- strictions.

Pair n° Relative length Arm ratio Type

ee 148.72 + 18.32 1.97 + 0.20 sm

135.58 = 18.76 1.71 = 0.63 sm.

3 126.80 + 10.50 4.69 + 0.48 st

4* 116.74 + 12.13 2.28 = 0.74 sm

5 O22 == 2227, 1.43 + 0.50 m

6* O39) = LAI 2.05 + 0.60 sm

7 74.03 + 8.03 2.10 = 0.32 sm

8 66.16 = 6.46 1.42 + 0.24 m

9 61.76 + 6.00 1.30 + 0.27 m

10 SI2D = O43 1.20 = 0.18 m 11 SIGS) 22 5.04! 1.90 + 0.15 sm 12 48.73 + 3.71 1.50 = 0.27 m 13 39)./D 22 Da) 1.40 + 0.12 m

grandisonae and Diaz & Nufiez (1988) re- ported Alsodes verrucosus (adult and tad- poles). The other locality where Alsodes kaweshkari was collected is Seno Huemu- les. This area has the same ecological char- acteristics as Puerto Edén. The male para- type, collected at a border of a cold stream, also has the nuptial asperities on fingers and chest.

Chromosomes.—Examination of 10 metaphase plates from the holotype re- vealed a diploid number of 2N = 26. All chromosomes are bi-armed and the funda- mental number (NF) is 52. Pairs 5, 8-10, 12, 13 are metacentric, pairs 1, 2, 4, 6, 7, 11 are submetacentric, and pair 3 is subtelo- centric. Pairs 1 and 4 have secondary con- strictions on the smaller arm, and pair 6 ex- hibits secondary constrictions on the longer arm. The karyotype of Alsodes kaweshkari is shown in Fig 5A. A summary of the rel- ative length, arm ratio, and type of chro- mosomes is presented in Table 2.

The C-banded karyotype, based on five plates (Fig 5B) shows constitutive hetero- chromatin in the pericentromeric region of all chromosomes. Thin heterochromatic

527

bands can be discerned at some telomeres, especially in those of pairs 1, 2, 6 and 10. A thin insterticial band was observed in the long arm of pair 5. Pair 4 presents a poly- morphic situation: one chromosome shows a remarkable band of pericentromeric po- sition, but in the other the band is located in the centromeric region. This situation was observed in all the examined plates.

Hemoglobin.—Figure 6A shows the chromatographic profiles of the hemoly- sates of Alsodes kaweshkari and A. monti- cola obtained from the fractions collected in the Sephadex G-50 minicolumn. The fractions with the higher absorbance values at 410 nm (fraction 9 in Alsodes kaweskari and fraction 8 in A. monticola) were elec- trophoretically analyzed under the experi- mental conditions used in this study. Figure 6B indicates the electrophoretical patterns obtained for these species. Both taxa share a similar two-banded pattern with an anodal band (Hb,) of 14.2 KDa an other catodal one (Hb,) of 16.8 KDa.

Etymology.—The new taxon is named for the Kaweshkar indians, a brave people, hunters of sea-lions, who still live in Puerto Edén, the type locality of the new species.

Comparisons

When the adult snout—vent length of AI- sodes kaweshkari (males, 59.9—62.2 mm.) is compared with that of the other Alsodes species, it is observed that this frog is as large as A. nodosus (58.5 mm), A. barrioi (59.9 mm) and A. tumultuosus (61.5 mm). Other members of the genus are small [A. vittatus (37.4 mm), A. gargola (40.8 mm), and A. verrucosus (42.4 mm)] and medium- sized [A. monticola (49.1 mm), A. montan- us (50.2 mm), A. vanzolinii (51.1 mm), and A. pehuenche (52.3 mm)]. Among the large-sized frogs, the external morphology of the foot provides useful characteristics to identify each taxon. One example of this is A. nodosus, a species characterized by the absence of lateral fringes (present on the toes of A. barrioi, A. kaweshkari, and A.

528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A B 0.6 KDa 1 Js 3 0.5 A. kaweshkar! —-29.0— uy E 9414 monticola = , O < GS a 20.0— = OFZ 0.1 ~—_ ** Hb2 14.2— Hb; O O 4 8 12 16 20 Fractions Fig. 6. Hemoglobins of Alsodes kaweshkari and A. monticola. (A) Chromatographic elution profiles of A.

monticola and A. kaweshkari in Sephadex G-50 minicolumn (B) Electrophoretical patterns of Alsodes kaweshkari and A. monticola. (1) Standard of molecular weight, (2) A. monticola, (3) Alsodes kaweshkari.

tumultuosus). In the case of this latter spe- cies, its toes are long and thin, whilst in Alsodes kaweshkari and A. tumultuosus they are thick and strongly fringed. These frogs also differ in characteristics of the outer metatarsal tubercle (large in Alsodes kaweshkari and small in A. tumultuosus). On the other hand, both species differ in webbing; reduced in Alsodes kaweshkari and moderately developed in A. tumultuo- sus (Fig. 7A, B).

Formas & Vera (1983) studied the kary- ological relationships among the member of the genus Alsodes. They recognized three groups within the genus: the barrioi group (2N = 34) (A. barrioi), the monticola group (2N = 26) (A. gargola, A. monticola, A. tumultuosus, A. vanzolinii, and A. verruco- sus), and the nodosus group (2N = 22) (A.

nodosus). The presence of 26 chromosomes in Alsodes kaweshkari allows its inclusion in the monticola group.

Muir (1981) studied the electrophoretical patterns of the hemoglobin molecule of 14 taxa of the frogs of the genus Xenopus. The species and subspecies were characterized and grouped according the number of bands obtained in the electrophoretical profiles. Xenopus borealis, X. fraseri, X. muelleri, X. ruwenzoriensis, X. tropicalis, X. vestitus and X. witteii show an electrophoretical pat- tern of two bands (Hb,, Hb,). Alsodes kaw- eshkari and A. monticola show a similar two-banded electrophoretical pattern like that found in some Xenopus species. Though our biochemical study was unable to differenciate Alsodes kaweshkari and A. monticola, this is the first attempt to char-

VOLUME 111, NUMBER 3 529

A

bas S = =

5mm

Fig. 7. Schematic plantar views of the left foot of Alsodes kaweshkari (A) and A. tumultuosus (B).

acterize the molecule of hemoglobin of de Chile (DBCG), Chile; Instituto de Biolo-

frogs of the genus Alsodes. gia Animal, Universidad Nacional de Cuyo (IBA), Argentina; Instituto de Zoologia, Specimens Examined Universidad Austral de Chile (IZUA), Chile;

Abbreviations.—Carmen Ubeda (personal Museo Nacional de Historia Natural collection) (CU), Argentina; Departamento (MNHN), Chile; Museo de Zoologia, Univ- de Biologia Celular y Genética, Universidad ersidad de Concepci6dn (MZUC), Chile.

530

Alsodes barrioi: YZUA 1629-1630; Cordil- lera Pelada, Provincia de Valdivia, 1020 m, Chile.

Alsodes gargola: CU 6; Macizo Loncoluan, Provincia de Neuquén, 1900 m, Argen- tina.

Alsodes montanus: IZUA 824; Estero Cob- arrubias, Provincia de Santiago, 2400 m, Chile.

Alsodes monticola: TZUA 1550,1749; Cor- dillera Pelada, Provincia de Valdivia, 1020 m, Chile.

Alsodes nodosus: YZUA 756,700; Aguas Claras, Provincia de Petorca, 150 m, Chile.

Alsodes pehuenche: IBA 1643; Valle del Pehuenche, Provincia de Mendoza, 2500 m, Argentina.

Alsodes tumultuosus: DBCG 161-162; La Parva, Provincia de Santiago, 2600 m, Chile.

Alsodes vanzolinii: MZUC 12063-12070; Ramadillas, Provincia de Arauco, 100 m, Chile.

Alsodes verrucosus: MNHN 1506; Puerto Edén, Provincia de Ultima Esperanza, 10 m, Chile.

Alsodes vittatus: MZUC (untagged); Cor- dillera de Pemehue, Provincia de Malle- co) 1152) Chile:

Acknowledgments

The authors would like to give special thanks to Pablo Corti who collected the paratype in Seno Huemules. Alberto Velo- so, Patricia Iturra, José Navarro (Universi- dad de Chile), Enrique Pereira (Universidad Nacional de Cuyo, Argentina) and Carmen Ubeda (Universidad del Comahue, Argen- tina) kindly provided specimens for com- parisons. Marcos Navarro expertly prepared the drawings. This work was supported by Direccion de Investigacion Universidad Austral de Chile, Proyecto S 95-25.

Literature Cited

Bogart, J. P. 1970. Systematic problems in the am- phibian family Leptodactylidae (Anura) as in-

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

dicated by karyological analysis —Cytogenetics 9:369—-383.

Cei, J. M. 1962. Batracios de Chile. Ediciones de la Universidad de Chile, Santiago, Chile, 128 pp.

. 1980. Amphibians of Argentina.—Monitore Zoologico Italiano (N.S.) Monografia 2:1—609.

Diaz, N., & H. Nunez. 1988. Nuevo hallazgo de Al- sodes verrucosus (Philippi 1902) en Chile y descripciédn de su larva.—Boletin Museo Na- cional Historia Natural, Chile 41:87—94.

di Castri, EF 1968. Esquisse écologique du Chili. Biol- ogie de l’Amerique australe. Cl. Debouteville and E. Rappaport, eds.—Editions du Centre Na- tional de la Recherche Scientifique, Paris 1V:7— 2.

Duellman, W. E. 1970. The hylid frogs of the Middle America.——Museum of Natural History, Uni- versity of Kansas Monographs 1:1—753.

Formas, J. R. 1989. Sinonimia e identidad de la rana austral Chilena Eupsophus vittatus (Philippi 1902) (Anura: Leptodactylidae).—Boletin So- ciedad de Biologia, Concepci6n 60:123—127.

. 1991. The karyotypes of the Chilean frogs

Eupsophus emiliopugini and E. vertebralis

(Amphibia: Anura: Leptodactylidae).—Pro-

ceedings of the Biological Society of Washing-

ton 104:7-11.

, & M. I. Vera. 1983. Karyological relation- ships among frogs of the genus Alsodes, with description of the karyotypes of A. vanzolinii and A. verrucosus.—Copeia 1983:1104—1107.

Frost, D. R. 1985. (ed). Amphibian species of the world. A taxonomic and geographic reference. Allen Press and the Association of Systematics Collection, Lawrence, Kansas, 732 pp.

Laemmli, U. K. 1970. Cleavage of structural proteins during the assambly of the head of bacterio- phage T,.—Nature 227:680—685.

Levan A., Fredga A., & A. Sandberg. 1964. Nomen- clature for centromeric positions on chromo- somes.—Hereditas 52:201—220.

Lynch, J. D. 1975. A new Chilean frog of the extra- andean assamblage of Telmatobius (Amphibia: Leptodactylidae).—Bulletin of Southern Cali- fornia Academy of Sciences 74:160—161.

Muir, A. V. 1981. Comparison of hemoglobins from the genus Xenopus (Amphibia: Salientia).— Journal of Experimental Zoology 218:327-—333.

Philippi, R. A. 1902. Suplemento de los Batraquios Chilenos descritos en la Historia Fisica y Poli- tica de Chile de don Claudio Gay. Santiago, 61 Pp.

Sumner, A. T. 1972. A simple technique for demon- strating centromeric heterochromatin.—Experi- mental Cell Research 75:304—306.

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111(3):531—534. 1998.

Helicoprion nevadensis (Wheeler, 1939) from the Pennsylvanian—Permian Antler Peak Limestone, Lander County, Nevada (Pisces: Selachii: Helicoprionidae)

Rex Alan Hanger and Ellen E. Strong

(RAH) Department of Geology, George Washington University, Washington, D.C. 20052, U.S.A.; (EES) Department of Biological Sciences, George Washington University, Washington, D.C. 20052, U.S.A.

Abstract.—A new specimen of Helicoprion nevadensis (Wheeler 1939) is described from the Antler Peak Limestone of Lander County, Nevada. The occurrence is significant as the only other individual of the species known has no geographic or stratigraphic information. The new specimen is dated as Wolf- campian (Early Permian) using associated fusulinids. Presence of H. nevadensis in Nevada and similar species in California confirms open-marine connections between the Permian shelf of North America and coeval island arcs to the west.

A new specimen of the spiral tooth-whorl shark fossil, Helicoprion nevadensis (Whee- ler 1939) is described from the Pennsylva- nian-Permian Antler Peak Limestone of Lander County, Nevada. This is only the second individual of the species discovered. The original geographic and stratigraphic lo- cation of the holotype are unknown. In con- trast, this specimen is precisely located and accurately dated for the first time. It is Wolf- campian (Lower Permian) based upon co- occurrence of fusulinids. Although this oc- currence does not resolve the controversy surrounding longitudinal separation between the North American continental margin and coeval island arcs to the west (Harwood and Miller 1990), this accurate locality and bio- Stratigraphic information does allows us to conclude that open marine connections ex- isted between North American (autochtho- nous) shelf seas and island arc (allochtho- nous) deposits during the Early Permian.

Systematic Paleontology Family Helicoprionidae Bendix-Almgreen, 1966 Genus Helicoprion Karpinsky, 1899 Helicoprion nevadensis Wheeler, 1939 Fig.) 1

Helicoprion nevadensis.—Wheeler, 1939: 109-112, fig. 3.

Description.—The symphysial whorl of the specimen consists of 1.5 volutions and is broken off at both juvenile and adult ends (Fig. 1). The maximum preserved diameter (tooth crown tip to tooth crown tip) is 100.35 mm. Description is compromised by the fact that the enclosing rock is broken so that most of the specimen is split laterally, and several tooth crowns are broken off at the shaft and missing. 47 tooth crowns are preserved in one volution. For a well-pre- served tooth at approximately 1 volution, the volution height (V) is 20.2 mm, the shaft height (S) is 2.3 mm, with an S/V ra- tio of 0.11. No serration denticles are pre- served.

Depository.—The specimen is deposited in the University of California Museum of Paleontology (UCMP) as specimen 140632.

Location.—From UCMP Locality V94012, in western Lander County, Neva- da, with a Township/Range coordinate of T32N, R44E, in NE % of SE % of Section 21 on the Snow Gulch, Nevada, 7.5-minute U.S. Geological Survey quadrangle, 1991 edition. The specimen occurs in pale brown

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Fig. 1. X 1.1.

siltite of the siliclastite-dominated facies of the Antler Peak Limestone (Theodore 1994).

Comparison.—Several workers (Bendix- Almgreen 1966, Siedlecki 1970) have noted the very strong similarities of all described Helicoprion species. Unfortunately, Heli- coprion fossils are always found as isolated specimens, not populations. The possibility exists that much of the published species diversity of the genus actually reflects the range of morphologic variation of very few, or even a single species. Because of this taxonomic imprecision and the incomplete- ness of UCMP 140632, specific taxonomic placement is difficult. For S/V ratio and number of tooth crowns/volution, UCMP 140632 is most like Wheeler’s (1939) ho- lotype H. nevadensis, UCMP 1001.

Helicoprion nevadensis. (UCMP 140632) from the Antler Peak Limestone, Lander County, Nevada,

Biostratigraphy.—UCMP Locality V94012 occurs between strata containing fusulinid foraminifera (Verville et al. 1986). Several species of Schwagerina confirm a Wolfcampian age for UCMP 140632, con- sistent with the established geologic range of Helicoprion (Wolfcampian—Guadalupi- an).

Biogeography.—The genus Helicoprion has a worldwide distribution, with species described from Russia (Karpinsky 1899, Obruchev 1953, for summary), Japan (Yabe 1903, Araki 1980), Laos (Hoffet 1933), Australia (Teichert 1940), Spitsbergen (Siedlecki 1970), Arctic Canada (Nassichuk & Spinosa 1970, Nassichuk 1971), British Columbia, Alberta (Logan & McGugan 1968), Idaho, Wyoming (Hay 1907, 1909; Williams & Dunkle 1948; Bendix-Alm-

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green 1966), Nevada, California (Wheeler 1939, Larson & Scott 1955, this report) West Texas (Kelly & Zangerl 1976, Chorn 1978), and Mexico (Mulleried 1940), Thus, Helicoprion is found within rocks of many Permian benthic faunal provinces (see for example, Shi et al. 1995, Yancey 1975, Bambach 1990). Collectively, these prov- inces span polar to equatorial paleolatitudes in both hemispheres, a remarkable distri- bution, even for a nektic organism, sug- gesting that temperature was not a limiting factor for Helicoprion. Paleogeography.—Documenting the oc- currence of Helicoprion is problematic in the paleogeographically “‘suspect’’ accreted terrane region of western North America. Besides UCMP 140632, only three individ- ual specimens are known. The holotype of H. nevadensis Wheeler, 1939, described originally as from the Koipato Group of the Humboldt Range, Nevada, is considered by Silberling (1973) not to have been obtained from the Koipato Group because of dis- crepancies between the lithology of the rock fragments attached to the fossil and the diagnostic lithologies of the Koipato Group. Helicoprion sierraensis Wheeler, 1939 is said to be found by an amateur “‘in a gla- cially transported boulder in the valley of Frazier Creek, California.’’ Convoluted in- terpretations have to be proposed to attempt to determine provenance for the fossil. A third reported specimen from Nevada, H. sp., is reasonably located, but comes from autochthonous rocks near Elko, Nevada (Larson & Scott 1955). This specimen is not figured in any publication, nor is its mu- seum deposition mentioned. The current lo- cation of the specimen is unknown. Heretofore, lack of adequate documen- tation for the genus has prevented reliable paleogeographic inference. The joint occur- rence of Helicoprion in autochthonous rocks of the Early Permian continental mar- gin (Wheeler 1939, Larson & Scott 1955, Roberts et al. 1958, this report) and in the allochthonous Northern Sierra terrane of McCloud Belt faunal affinity (Harwood

533

1992, Miller 1987) should not be taken as evidence of original paleogeographic prox- imity of the McCloud Belt to North Amer- ica during the Permian, nor can it resolve the problem of longitudinal separation. However, presence of the nektic genus in both areas does suggest presence of open marine conditions between these tectonic elements.

Acknowledgments

R. T. Ashinhurst of the Battle Mountain Gold Company, Battle Mountain, Nevada kindly donated the specimen to UCMP.

Literature Cited

Araki, H. 1980. The discovery of the fossil genus Hel- icoprion, a chondrichthian, from Kesennuma City, Miyagi Prefecture, Japan.—Geological Society of Japan, Journal 86:135—-137.

Bambach, R. K. 1990. Late Paleozoic provinciality in the marine realm. Pp. 307-323 in W. S. Mc- Kerrow and C. R. Scotese, eds., Palaeozoic Pa- laeogeography and Biogeography. Geological Society of London Memoir Number 12, Lon- don, 431 pp.

Chorn, J. 1978. Helicoprion (Elasmobranchii, Edesti- dae) from the Bone Spring Formation (Lower Permian) of West Texas.—University of Kansas Paleontological Contributions, Paper 89:2—4.

Bendix-Almgreen, S. E. 1966. New investigations on Helicoprion from the Phosphoria Formation of South-East Idaho, U.S.A.—Biologiske Skrifter Danske Videnskabermes Selskab 14(5):1—54.

Harwood, D. S. 1992. Stratigraphy of Paleozoic and Lower Mesozoic rocks in the Northern Sierra terrane, California.—United States Geological Survey Bulletin 1957:1—78.

, & M. M. Miller (eds.). 1990. Paleozoic and Early Mesozoic Paleogeographic Relations; Si- erra Nevada, Klamath Mountains, and Related Terranes. Geological Society of America, Spe- cial Paper 255, Boulder, Colorado, 422 pp.

Hay, O. P. 1907. A new genus and species of fossil shark related to Edestus Leidy.—Science, New Series 26(653):22—24.

. 1909. On the nature of Edestus and related genera, with descriptions of one new genus and three new species.—Proceedings of the U.S. National Museum 37:43—61.

Hoffet, J. H. 1933. Etudes géologiques sur le contre de Il’Indochine entre Tourane et le Mékong.— Bulletin, Service Géologique de 1’Indochine 20(2):1-154.

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Karpinsky, A. 1899. Uber die Reste von Edestiden und die neue Gattung Helicoprion.—Verhandlung Russisch-Keiserling Mineralogische Gesells- chaft zu St. Petersburg Series 2, 36(II]):361—376.

Kelly, M. A., & R. Zangerl. 1976. Helicoprion (Edes- tidae) in the Permian of West Texas.—Journal of Paleontology 50:992—994.

Larson, E. R., & J. B. Scott. 1955. Helicoprion from Elko County, Nevada—Journal of Paleontolo- gy 29:918-919.

Logan, A., & A. McGugan. 1968. Biostratigraphy and faunas of the Permian Ishbel Group, Canadian Rocky Mountains.—Journal of Paleontology 42:1123-1139.

Miller, M. M. 1987. Dispersed remenants of a north- east Pacific fringing arc: Upper Paleozoic island arc terranes of McCloud Belt faunal affinity, western United States.—Tectonics 6:807—830.

Mullereid, EF K. G. 1945. El edestido, Helicoprion, en contrado por primera vez en Mexico, en el es- tado de Coahuila.—Ciencia Revista Hispano- America de Ciencias Puras y Aplicadas 6:208— 211.

Nassichuk, W. W. 1971. Helicoprion and Physonemus, Permian vertebrates from the Assistance For- mation, Canadian Arctic Archipelago—Geo- logical Survey of Canada, Bulletin 192:83—93.

, & C. Spinosa. 1970. Helicoprion sp., A Perm- ian elasmobranch from Ellesmere Island, Ca- nadian Arctic.—Journal of Paleontology 44: 1130-1132.

Obruchev, D. V. 1953. A study of the edestids and the work of A. P. Karpinsky.—Akademia Nauk SSSR, Trudy Paleontologischkogo Instituto 45: 1-88.

Roberts, R. J., PR E. Hotz, J. Gilluly, & H. G. Ferguson. 1958. Paleozoic rocks of North-Central Neva- da.—American Association of Petroleum Ge- ologists Bulletin, 42:2813—2857.

Shi, G. R., N. W. Archbold, & L.-P. Zhan. 1995. Dis-

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tribution and characteristics of mixed (transi- tional) mid-Permian (Late Artinskian—Ufimian) marine faunas in Asia and their palaeogeo- graphical implications.—Palaeogeography, Pa- laeoclimatology, Palaeoecology 114:241—271.

Siedlecki, S. 1970. A Helicoprion from the Permian of Spitsbergen.—Norsk Polarinstitutt Arbok 1968:36—54.

Silberling, N. J. 1973. Geologic events during Perm- ian-Triassic time along the Pacific margin of the United States, Pp. 345-362 in A. Logan and L. V. Hillis, eds., The Permian and Triassic sys- tems and their mutual boundary. Canadian So- ciety of Petroleum Geologists, Memoir 2. Cal- gary, Alberta., 766 pp.

Teichert, C. 1940. Helicoprion in the Permian of West- ern Australia—Journal of Paleontology 14: 140-149.

Theodore, T. G. 1994. Preliminary geologic map of the Snow Gulch quadrangle, Humboldt and Lander Counties, Nevada. United States Geological Survey, Open-file Report 94-436.

Verville, G. J.. G. A. Sanderson, & D. D. Drowley. 1986. Wolfcampian fusulinids from the Antler Peak Limestone, Battle Mountain, Lander County, Nevada.—Journal of Foraminiferal Re- search 16:353-—362.

Wheeler, H. E. 1939. Helicoprion in the Anthracolithic (Late Paleozoic) of Nevada and California, and its stratigraphic significance.—Journal of Pale- ontology 13:103—114.

Williams, J. S., & D. H. Dunkle. 1948. Helicoprion- like fossils in the Phosphoria Formation.—Geo- logical Society of America, Bulletin 59:1362.

Yabe, H. 1903. On a Fusulina-Limestone with Heli- coprion in Japan.—Geological Society of To- kyo, Journal 10:1—13.

Yancey, T. E. 1975. Permian marine biotic provinces in North America.—Journal of Paleontology 49: 758-766.

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111(3):535-—550. 1998.

Pentamera rigida and P. pediparva, two new species of sea cucumber from the west coast of North America (Echinodermata: Holothuroidea)

Philip Lambert

Natural History Section, Royal British Columbia Museum, P.O. Box 9815 Stn Prov Govt, Victoria, B.C., Canada V8W 9W2

Abstract.—Two new species of the genus Pentamera from the west coast of North America are described, and all known species from the west coast of North and South America are reviewed. Pentamera rigida is a slender, stiff, U-shaped species with five rows of podia. It occurs in sand-gravel substrata from 18 to 421 m between British Columbia and California. The table ossicles of the skin have an angular disc with a broad, low, convoluted spire. Pentamera pediparva has a soft, curved body with five rows of fine podia. It is known from central British Columbia to northern California from 8 to 120 m in mud- gravel or sand-gravel. The table ossicles are small and oval with small, two pillared spires. The taxonomic characters of fourteen species of Pentamera are

summarized.

Twelve species of Pentamera have been described from the west coast of North and South America (Stimpson 1851, 1864; Lud- wig 1886a, 1886b; Clark 1924; Deichmann 1938a, 1938b). Panning (1949) placed the genus Pentamera in Thyoninae, a new sub- family of Cucumariidae. Based on the pres- ence of posterior processes composed of a mosaic of small pieces, Pawson & Fell (1965) transferred Thyoninae Panning to the family Phyllophoridae. No new species of Pentamera have been added to the west coast fauna since Deichmann (1938a, 1938b), with the exception of those trans- ferred to the genus by Panning (1949). The purpose of this paper is to describe two new species of Pentamera from the Pacific coast of North America and present a summary table of morphological characters for spe- cies from the west coast of North and South America.

Materials and Methods

Ossicle slides were prepared by the method described in Lambert (1985). At

least 30 ossicles per specimen were mea- sured on transects across each slide. Length and width of disc, and width of central spire were recorded for tables. Length, height of curve, and height of spire were recorded for supporting tables (Fig. 7). Only ossicles that were totally within the field of vision, lying flat, and not broken, were measured. Via a drawing tube attached to a Wild M20 microscope, the image of each ossicle was visualised on a digitising tablet (Summas- ketch II, Summagraphics Corporation). Measurements were made with the cursor and automatically saved to a file using SigmaScan software (Jandel Scientific). That file was then transferred to a statistical package (Systat, Systat, Inc.) for analysis. External measurements of the whole spec- imen were made with an ocular micrometer, calipers, or in the case of strongly curved specimens, a string was used to trace the distance from the base of the tentacles to the anus, around the outer side of the curve, usually the ventral side.

Institutions mentioned in this paper are abbreviated as follows: Allan Hancock

536

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Fig. 1. length 7 cm, RBCM 984-199-3.

Foundation (AHF) now housed at LACM; California Academy of Sciences, San Fran- cisco (CASIZ); Fisheries Research Board of Canada (FRB) now known as Department of Fisheries and Oceans, (DFO); Canadian Museum of Nature, Ottawa (CMNI or NMC); Natural History Museum of Los Angeles County (LACM); Royal British Columbia Museum, Victoria (RBCM); Na- tional Museum of Natural History (USNM), Washington, D.C.

Results Order Dendrochirotida Grube Family Phyllophoridae Oestergren

Diagnosis.—Dendrochirotid holothurians without a test. Tentacles 10—25 (usually 10 or 20), well branched. Pedicels either re- stricted to the ambulacra or scattered, but fewer and usually papilliform dorsally. Cal- careous ring complex, often tubular, with long or short posterior processes, both ring and processes always composed of a mo- saic of small pieces. Ossicles usually plates or buttons, rods, tables or derivatives of ta- bles (after Thandar 1990).

Remarks.—Thandar modified the diag- nosis of Pawson (1982) by adding that the podia may be in rows or scattered and that

Preserved specimen of Pentamera rigida dredged from type locality, 115 m, off Nootka Sound, B.C.,

the ossicles may be tables or derivatives of tables in the form of plates, rods or buttons.

Subfamily Thyoninae Panning, 1949

Diagnosis.—Small to medium-sized phyllophorids, rarely more than 100 mm long. Body soft. Pedicels numerous, usually scattered all round, but often most crowded ventrally. Tentacles 10, ventral two always reduced. Calcareous ring tubular with long paired posterior processes on radial plates. Ossicles of body wall usually in the form of tables or plates (buttons), or absent (after Thandar 1990).

Genus Pentamera Ayres, 1852

Diagnosis.—Small to medium sized forms. Ten tentacles with two smaller ven- tral tentacles. Nonretractile podia in five se- ries, not scattered. Radials of the calcareous ring with long forked tails. In the skin two pillared tables or derivatives of these with the spire reduced or absent. Podia with large endplates and curved supporting ta- bles with spires varying from low to high. Tentacles usually with rods, plates or both. (After Deichmann 1941)

Type species: Pentamera pulcherrima Ayres

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33)

Fig. 2. introvert. (C) Tentacle ossicles. (D) Supporting tables and end plates of podia. (E) Calcareous ring and retractor muscles; scale bar 5 mm. All from holotype (RBCM 997-123-1) except for the podial end plates (RBCM 983- 1658-30) and the calcareous ring of paratype (RBCM 984-199-3). Top scale bar 100 ym applies to all ossicles.

Pentamera rigida, new species Figs. 1-3

Pentamera lissoplaca.—Bergen 1996 (par- tim):239, fig. 9.23B. Specimens de- scribed as having “‘large tables’’ may be P. rigida.

Pentamera sp. A Lambert, 1997:101, figs. 52-53, photo 24.

Diagnosis.—Body U-shaped; skin usu-

Ossicles and calcareous ring of Pentamera rigida. (A) Tables of the dorsal skin. (B) Plates from the

ally stiff and white. Podia in five series, longer and more abundant in midventral re- gion than distally. Ten small dendritic ten- tacles including smaller ventral pair; often speckled with brown. Calcareous ring long and tubular with forked tails on the radials; made up of a mosaic of smaller pieces; ratio of radial to interradial length approximately 4:1. Skin ossicles circular, triangular or star- shaped tables (100-300 sm) with a broad

538

Fig. 3.

Collection sites for Pentamera rigida.

convoluted spire covering about one-half width of disk; small, curved supporting ta- bles (100 pm) with low spire. Introvert with oval knobbed perforated plates, some with remnant of a spire. Tentacle ossicles in two forms, finer curved oval plates and large ro- bust rods with a few holes.

Material examined.—Eighty-one speci- mens from 15 localities in British Colum- bia, one locality in Oregon, four in Califor- nia, and one from an unknown locality on the west coast of U.S.A. Number of speci- mens in parentheses after the catalogue number; depth in metres (m).

Holotype.—RBCM 997-123-1 (1), col-

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lected by E Bernard and T. C. Lambert, FRB station 63-91, 8 Sep 1964, length 6.8 cm (measured along the outside of the curved body), female.

Type locality.—British Columbia, Van- couver Island, Nootka Sound, 49°32'N, 127°03'W, 115 m, sandy mud.

Paratypes.—RBCM 984-248-1 (1), col- lected by D. B. Quayle, FRB Haul No. 1, 7 Jul 1962, British Columbia, Kitkatla Inlet, Porcher Island, 53°51'N, 130°W, 37 m, sand and gravel, specimen 10.8 cm long. RBCM 984-200-1 (7), FRB station 63-204, 10 Sep 1964, British Columbia, Vancouver Island, Quatsino Sound, 50°19.6’N, 128°06.5'W, 119 m, sand, specimen lengths from 2.8 to 5.1 cm. RBCM 984-199-3 (12), collected by E Bernard and T. C. Lambert, FRB sta- tion 63-91, 8 Sep 1964, British Columbia, Vancouver Island, Nootka Sound, 49°32’N, 127°03'W, 115 m, sandy mud, from same lot as holotype, specimen lengths from 3 to 8 cm (mean 5.5). USNM E16915 (32), col- lected by A. Carey, Oregon State Univer- sity, Cruise C 700810, BMT 239, 17 Aug 1970, Oregon, off Hecata Head Light, 44°08.8'N, 124°24'W, 100 m, specimen lengths from 1.5 to 5.7 cm. CASIZ 50227 (1), Sta. 20, 28 Sep 1949, California, off Point Reyes, south of Cordell Bank, 37°55.75'N; 123°19:25'W,. PLO= ise specimen 6 cm long. AHF 440.61 (1), Ve- lero 6131-59, California, Ventura Co., Ox- nard, Port Hueneme Lighthouse, 34°8.1'N, 119°21.5'W, 165 m.

Other collections.—British Columbia: RBCM 989-564-4 (1), collected by D. B. Quayle, FRB station 70-24, Jul 1970, Bramham Island near Pine Island, 51°05’N, 27°39'W, 82 m. RBCM 984-213-1 (1), col- lected by D. B. Quayle, FRB station 71-14, Apr 1971, Gordon Channel, near Nigei Is- land, 50°53. 1'N,: 127°36.3' W942 See RBCM 990-939-11 (2), FRB station 67-66, Aug 1967, Vancouver Island, Cape Scott, 50°39.8'N, 128°47.2'’W, 200 m, gravel. RBCM 984-249-1 (1), collected by J. Fleu- ry, FRB Drag #7 + #8, 27 Jun 1962, Baker Pass, near Cortes Island, 50°04.0’N,

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Fig. 4. Holotype of Pentamera pediparva collected by scuba from 7.5 m in Quatsino Sound, B.C., length 8

cm, RBCM 997-124-1.

124°59.0’W, 117 m, gravel. RBCM 980- 344-1 (1), collected by P. Lambert, station L80-61, 6 Jul 1980, Vancouver Island, Kyu- quot Channel, Sandy Bay, Rugged Point, 49°58.2’N, 127°14.6'W, 9 m, clean sand and algae. RBCM 55-56 (1), collected by D. El- lis, 16 Aug 1965, Strait of Georgia, 49°54.05'N, 125°04’W, 178 m, sand. RBCM 983-1658-30 (3), collected by Gor- don Green, 23 Nov 1983, Vancouver Island, off Kyuquot Sound, 49°45.7'N, 127°30.1’W, 150 m, sand. RBCM 974-570- 6 (3), FRB station 2231-33, 17 Jul 1934, Vancouver Island, Estevan Point, 49°22.3'’N, 126°55'W, 137 m, sand. RBCM 986-93-27 (3), collected by P. Lambert, on Endeavour, Station L86-9, 19 Mar 1986, Vancouver Island, Estevan Point, 49°11'N, 126°45.2'W, 120 m, sand. RBCM 983- 1397-1 (1), collected by M. Byrne, 11 Feb 1983, east of Galiano Island, 49°01.48'N, 123°29.45'W, 50—230 m. RBCM 991-10-1 (2), collected by S. Carson, 22 Feb 1990, Barkley Sound, Trevor Channel, 48°52'N, 125°08'W, 54 m, mud. RBCM 988-758-10 (1), collected by P. Lambert, G. Green, D. Bright, Station L88-27, 22 Jun 1988, Sat- ellite Channel, Boatswain Bank, 48°42.2'N, 123°32.1'W, 18 m.

California: CASIZ uncatalogued (1), 9 Aug 1940, 2.5 mi. off Mad River, Eureka,

37 m. USNM E2371 (4), Albatross St. 2902, California, Channel Islands, Santa Cruz Channel, Santa Rosa Island, 34°6.0'N, 120°2.0'W, 97 m. USNM 30563 (1), Alba- tross St. unknown, West Coast United States, depth unknown.

Description.—Total length 1.5-10.5 cm (measured along the outer curve of the body); mean length (cm + 1 SD) 4.6 + 1.6 cm (n = 72). Holotype 6.8 cm long. Body typically bent in a tight U-shape with long slender anterior and posterior ends (Fig. 1). Skin stiff with ossicles. Podia slender and probably non-retractile because of their heavy complement of ossicles. Podia form five bands, each consisting of two rows, crowed together in the middle of the body on ventral side but sparse distally and on dorsal side. Specimens typically white in life and in alcohol. Ten tentacles arranged in five pairs with a smaller ventral pair.

Madreporite in dorsal mesentery about two-thirds of distance from anterior end of calcareous ring. Semi-circular madreporite with a long narrow stone canal, connects to the ring canal near posterior tip of calcar- eous ring. One polian vesicle usually on ventral side of ring.

Two respiratory trees emerge from cloa- ca on left and right sides; each one splits into a dorsal and ventral branch; dorsal

Fig. 5. the introvert. (C) End plate of podia (D) Tentacle ossicles. (E) Supporting tables of podia (F) Calcareous ring; scale bar, 5 mm. All from holotype (RBCM 997-124-1). Top scale bar (100 tm) applies to all ossicles.

branch runs full length of body, ventral branch only to mid body. Respiratory trees usually brown, with a main trunk and sim- ple, short side branches. Cloaca occupies most of narrow ‘“‘tail.”’

Gonad has two tufts of unbranched tu- bules, one tuft on each side of dorsal mes- entery where it joins the dorsal body wall.

Thin retractor muscles attach to body wall at a point level with posterior end of calcareous ring. Five thin longitudinal mus- cles. Radial plates of calcareous ring with long posterior prolongations; interradials with no tails, shaped like an elongate tri- angle (Fig. 2E). Transverse thickenings in- dicate joints between mosaic of pieces that make up the tails. Ratio of interradial to ra- dial length about 1:4.

Tables of mid skin circular to roughly tri-

See °

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Ossicles and calcareous ring of Pentamera pediparva. (A) Tables of the dorsal skin. (B) Plates from

angular or star-shaped, with a mean diam- eter (um + 1 SD) of 194 + 45 (n = 141) with a complex central spire (width 102 + 32, n = 140) that covers about one-half the surface area of table (Fig. 2A). Supporting tables of podia small (length 110 + 15,n = 90) and tightly curved with a low spire (height 16 + 5, n = 90) with three or four teeth. Ratio of length (L) to total height (TH) of supporting tables 1.9 + 0.3, n = 90 (see Fig. 7 for explanation of dimen- sions). Podial end plates small (diameter 115-135 pm, n = 5) (Fig. 2D). Ossicles in tail region not significantly different from mid skin. In juveniles (<2 cm) tables tend to be more star-shaped and spires not as ro- bust. Introvert has oval perforated plates with two larger central holes and knobs on the surface of plate (Fig. 2B). In tentacles,

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Fig. 6. Collection sites for Pentamera pediparva.

ossicles range in size from large flat rods (ength 230 wm) with a few knobs and holes to smaller curved perforated plates with scalloped edges (Fig. 2C). Etymology.—The species name, rigida, is based on the Latin word rigidus, meaning stiff or inflexible and refers to the typically stiff ossicle-filled body of this species. Distribution and habitat.—Pentamera rigida is known from Porcher Island near Prince Rupert, British Columbia (53°51'N, 130°W) south to Santa Cruz Channel, Santa Rosa Island, Channel Islands, California (34°6.0'N, 120°2.0’W) (Fig. 3). It ranges in depth from 18 to 421 m on the continental

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shelf. The majority of collections are from less than 200 m. The one specimen from 421 m is from a deep trench between Van- couver Island and the mainland. So far none have been collected on the continental slope or deeper. Usually occurs in sand or gravel substrata.

Pentamera pediparva, new species Figs. 4-6

Pentamera sp. B Lambert, 1997:104, figs. 54, 55, photo 25.

Diagnosis.—Body curved; up to 7 cm long; skin usually soft; colour white to

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Table