Skip to main content

Full text of "Transactions of the San Diego Society of Natural History"

See other formats


•wwV'^ v lw j^^^ TCTr' ;j 


, ,-»-..^.v^..s^>fti'.r?it,Tyrtf:rris^u^-.ia^..JMa 


Library of the 

Museum of 

Comparative Zoology 






1. Late Wisconsinan and Holocene fauna from Smith Creek Canyon, Snake 
Range. Nevada. By Jim I. Mead. Robert S. Thompson and Thomas R. 

Van De\ender. 6 August 1982 1-26 

2. Fossil decapod crustaceans from the Lower Cretaceous. Glen Rose Limestone 

of Central Texas. By Gale A. Bishop, 25 January 1983 27-55 

3. A new subspecies of Euphyes vestris (Boisduval) from southern California 
(Lepidoptera: Hesperiidae). By John W. Brown and William W. McGuire. 

24 June 1983 ' 57-68 

4. Two new iodeid isopods from Baja California and the Gulf of California 
(Mexico) and an analysis of the evolutionary history of the genus Colidotea 
(Crustacea: Isopoda: Idoteidae). By Richard C. Brusca. 24 June 1983 .... 69-79 

5. Descriptions of five new muricacean gastropods and comments on two 
additional species, in the Families Muricidae and Coralliophilidae: 
(Mollusca). By Anthony D'Attilio and Barbara W. Myers, 18 January 

1984 '. 81-94 

6. The fossil Leptostracan Rhahdouraea ^?A7/r; (Malzahn. 1958). By Frederick 

R. Schram and Eric Mabahn. 18 January 1984 95-98 

7. Phylogen\, evolution and biogeography of the marine isopod Subfamily 
Idoteinae (Crustacea: Isopoda: Idoteidae). By Richard C. Brusca, 18 
January 1984 99-134 

8. Rhamdia reddelli, new species, the first blind pimelodid catfish from Middle 
America, with a key to the Mexican species. B\ Robert Rush Miller. 18 
January 1984 '. ' 135-144 

9. A complete specimen of Peachella brevispina Palmer — an unusual olenellid 
trilobite (Arthropoda: Olenellida) from the lower Cambrian of California. 

By James H. Stitt and R. L. Clark. 20 June 1984 145-150 

10. Type specimens of amphibians and reptiles in the San Diego Natural History 
Museum. By Gregory K. Pregill and James E. Berrian. 20 June 1984 151-164 

11. Imocaris tuhenulata, n. gen., n. sp. (Crustacea: Decapoda) from the upper 
Mississippian Imo Formation. Arkansas. By Frederick R. Schram and 

Royal H. Mapes. 20 June 1984 165-168 

12. New material of Hydrodamalis custae (Mammalia: Dugongidae) from the 
Miocene and Pliocene of San Diego County, California. By Daryl P. 
Domning and Thomas A. Demere, 20 November 1984 169-188 

13. Fossil Syncarida. By Frederick R. Schram, 20 November 1984 189-246 

14. The late Wisconsinan vertebrate fauna from Deadman Cave, southern 
Arizona. Bv Jim 1. Mead. Edward L. Roth. Thomas R. Van De\enderand 

Da\id W. Steadman. 20 November 1984 247-276 

15. A Pliocene flora from Chula Vista, San Diego County, California. By 

Daniel I. A.xclrod and J homas A. Demere, 20 November 1984 277-300 

16. Relationships within Eumalacostracan Crustacea. By Frederick R. Schram. 

20 Noxcmber 1984 301-312 

17. Historv and status of the avifauna of Isla Guadalupe. Mexico. By Joseph R. 

Jehl, Jr. and William I. Everett, 30 January 1985 313-336 



Volume 20 Number 1 pp. 1-26 6 August 1982 

Late Wisconsinan and Holocene Fauna from Smith Creek Canyon, 
Snake Range, Nevada 

Jim I. Mead, Robert S. Thompson, and Thomas R. Van Devender 

Laboratory of Paleoenvironmental Studies, Department of Geosciences, 
University of Arizona, Tucson 85721 

Abstract. During the late Pleistocene, montane glaciers in the Snake Range, eastern Nevada 
reached an elevation as low as 2900 m and pluvial Lake Bonneville rose to approximately 1580 m. 
only 130 m below the entrance of the east-facing Smith Creek Canyon. It is not known whether the 
two events coincided. Packrat midden macrofossils indicate that bristlecone pine (Piiuis lont^'acva). 
limber pine (P. flexilis), and other subalpine taxa dominated the plant communities in Smith Creek 
Canyon through the late Wisconsinan. We report here 2 fish, 4 anurans, 9 lizards, 8 snakes, and 15 
small mammals recovered from 15 packrat middens and a pollen profile from cave fill. This assemblage 
adds 15 amphibians and reptiles and 7 mammals to the approximately 46 terrestrial animals previously 
known from the late Pleistocene and early Holocene of the canyon. Dung pellets of the locally extir- 
pated pika (Ochotona cf. princeps) were found in five packrat middens. A single tooth of the heather 
vole (Phenacomys cf. intermedius) from Smith Creek Cave is the first late Pleistocene record for this 
genus in the Great Basin. We review and update the late Pleistocene and Holocene fauna from 4 caves 
and 2 shelters in Smith Creek Canyon. 


The Snake Range of White Pine County, eastern Nevada, is a north-south trending 
mountain 80 km long. Smith Creek Canyon, a deep canyon on the eastern face of this 
Great Basin range, opens onto the Lake Bonneville playa in the Snake Valley of Utah. 
During the late Pleistocene, montane glaciers reached elevations as low as 2900 m in 
the Snake Range (Drewes 1958), and pluvial Lake Bonneville rose to a level of ap- 
proximately 1580 m, only 130 m below the entrance of Smith Creek Canyon. If the 
late Wisconsinan glacial maximum coincided with the high stand of Lake Bonneville, 
biotic communities in this canyon would have been restricted to an elevational range 
of less than 1310 m (Fig. I). 

Fossil localities. — Late Pleistocene vertebrate fossils from Smith Creek Canyon, 
specifically Smith Creek Cave, were reported by M. Harrington { 1934) from the South- 
west Museum and by others since. In 1955, T. E. Downs and associates from the 
Natural History Museum of Los Angeles County (LACM), California, screened some 
of the cave sediments. Field notes of the LACM indicate that the cave contained little 
or no stratification (Brattstrom 1976). Howard (1935, 1952) described a new species of 
extinct eagle (Spizaetiis willctti) and a teratorn (Teratornis incrcilihilis) and listed the 
Smith Creek Cave avifauna, unfortunately giving the stratigraphic associations only as 
from the older deposits of the lower levels. Stock (1936) described a new species of 
extinct mountain goat (Oreamnos harringtoni), but again did not discuss placement 
within the stratigraphic sequence. Brattstrom (1958, 1976) reported amphibians and 
reptiles from the cave sediments and Goodrich (1965) updated the list of the entire 
fauna (amphibians, reptiles, birds, and mammals); again the stratigraphy and temporal 
associations were only scantily discussed. 

Bryan ( I979«) made further excavations in the cave in search of evidence for Early 
Man and for the first time described the stratigraphy within a portion of the cave. 

Figure 1 . Map of Nevada and bordering states showing Smith Creek Canyon ( I ) and Gatecliff Shelter (2) 
along with Great Salt Lake (black) and Lake Bonneville (stippled) at the 1580 m elevation. 

Miller (1979) identified the mammalian fauna recovered during Bryan's excavations, 
providing another updated listing of the fauna; some stratigraphic associations were 
given. Both Bryan {\979a) and Miller (1979) emphasized the deposits and fauna asso- 
ciated with Man, those units dating less than 11500 B.P. Thus no comprehensive 
stratigraphic analysis of the fauna has been published. 

In 1977 and 1978 we visited Smith Creek Canyon to study fossil packrat (Neotoma) 
middens and to collect a pollen profile from the Smith Creek Cave sediments. Plant 
remains from some of these middens are discussed in a previous report (Thompson 
1979), and other assemblages are still being analyzed (RST). The packrat midden fossils 
document that bristlecone pine {Pinus longaeva), limber pine {P. fle.xilis), and other 
subalpine taxa dominated the plant communities in Smith Creek Canyon throughout 
the late Wisconsinan. The Smith Creek Cave pollen profile provided little paleoenvi- 
ronmental information and could not be dated directly. In this report we describe the 
fish, amphibians, reptiles, and mammals recovered from 15 packrat middens from 
Smith Creek Canyon and in association with the pollen profile, and the herpetofauna 
recovered by Bryan (\919a, Miller 1979) in his excavation in Smith Creek Cave. We 
also review the entire local fauna recovered from all archaeological and paleontological 

investigations in Smith Creek Canyon and attempt to place these records in the correct 
paleoenvironmental and chronological context. 

Local setting. — The lower concourse of Smith Creek Canyon is a steep-walled 
east-west trending canyon incised into Paleozoic limestone. In this part of the canyon 
there are numerous caves, including those described below in which we have found 
our fossil materials. The upper reaches of Smith Creek Canyon are lined with other 
Paleozoic sedimentary rocks and Mesozoic intrusives. We have not located any caves 
or fossil packrat middens in this area. The intermittent waters of Smith Creek are 
primarily derived from the elevated plateau surrounding Mount Moriah (3673 m). 

Although the vegetation in Smith Creek Canyon is broadly divided into elevational 
zones, the major slope and aspect contrasts between the north and south facing slopes 
create numerous microenvironments for plants of higher and lower elevations. At the 
canyon entrance the plant community is dominated by shadscale (A triplex confertifolia) 
and other xerophytes, including spiny hopsage {Grayia spinosa), greasewood (Sarcoh- 
atus venniciilatiis), rabbitbush {Chrysothamniis nanseosns). Mormon tea (Ephedra 
nevadensis), horsebrush (Tetradyinia axillaris), Harriman yucca (Yucca harriinaniae), 
and bud-sage (Artemisia spinescens). These same taxa are dominant in the Snake 
Valley to the east and also are common on xeric slopes throughout Smith Creek Can- 

Sagebrush (Artemisia tridentata, A. nova) occurs both in nearly pure stands in the 
canyon bottom and as a common understory element in the pinyon-juniper woodlands 
and forested communities. Utah juniper (Juniperus osteosperma) and single needle 
pinyon (Finns monophylla) occur throughout the limestone walled part of the canyon 
and form denser stands on the north facing slopes and on alluvium. Little leaf mountain 
mahogany (Cercocarpus intricatus), greasebush (Forsellesia nevadensis), skunkbush 
(Rhus trilohata), and joint-fir (Ephedra viridis) are common in the pinyon-juniper 
woodland and are often dominant on xeric slopes and on limestone substrates. 

Montane and subalpine conifers, including white fir (Abies concolor), Douglas fir 
(Pseudotsuga menziesii), Engelmann spruce (Picea engelmannii), ponderosa pine 
(Pinus ponderosa), limber pine, and bristlecone pine, are present at relatively low 
elevations in mesic niches on the north facing slope and along the upper concourse of 
Smith Creek. Other common riparian plants include narrowleaf cottonwood (Populus 
angustifolia), willows (Salix spp.), water birch (Betula occidentalis), chokecherry 
(Pruniis virginiana). Rocky Mountain maple (Acer glabrum). Rocky Mountain juniper 
(Juniperus scopulorum), and wild rose (Rosa woodsii). The higher elevations of the 
northern Snake Range support groves of quaking aspen (Populus tremuloides) and 
stands of bristlecone pine and limber pine on rocky outcrops. 

Although the fossil localities we investigated occur in a narrow elevational range 
(1860 m to 2060 m), the variations in slope and aspect at these sites place them in 
different environmental settings (Table 1). Smith Creek Cave (1950 m elev.) and Ladder 
Cave (2060 m elev.) are on a steep south facing slope at the entrance to the canyon 
and are surrounded by a xeric pinyon-juniper woodland with an abundance of xero- 
phytic plants (see Thompson 1979). Streamview Shelter is on a protected north facing 
slope near the canyon bottom (1860 m elev.), and Council Hall Cave is higher (2040 
m elev.) on the same slope. These two sites are in a relatively dense pinyon-juniper 
woodland, with fewer xerophytes than on the opposing slope. Montane and subalpine 
conifers occur in protected niches near both Council Hall Cave and Streamview Shel- 
ter. Amy's Shelter and the Kachina Cave are in the bottom of the canyon by the creek. 
These two sites probably were buried by alluvium during the last full-glacial, only to 
be exposed during the latest phases of the late glacial or early Holocene. 


The Smith Creek Canyon localities included within this report are Smith Creek, 
Ladder, Council Hall, and Kachina caves. Amy's Shcilcr and the packrat midden 
rockshelter we named Streamview. 

Table i. Paleontological and archaeological sites in Smith Creek Canyon, Snake Range, and the adjacent 
Snake Valley, Nevada, that are discussed in text. 




Type of deposit 

Age range 

Present status 

Smith Creek 


1950 m 

and non- 

sediments; 4 

^12 500 to 1000 

Relatively xeric site, sparse 
pinyon-juniper woodland, 
active talus formation, high 
above canyon bottom 

Ladder Cave 


2060 m 

6 packrat 

27 000 to 1 1 000 

Relatively xeric site, sparse 
pinyon-juniper woodland, 
active talus formation, high 
above canyon bottom 



1740 m 

cave sediments 

5000 to 1500 

In canyon bottom near 
entrance, relatively arid, 
scattered juniper with 
sagebrush-shadscale shrub 



1860 m 

3 packrat 

17 000 to 6500 

Ca. 15 m above canyon 
bottom, pinyon-juniper 

Kachina Cave 


1770 m 

cave sediments 

ca. 4500 to present 

Along streamside of 

intermittent Smith Creek, 
pinyon-juniper and riparian 

Council Hall 


2040 m 

Cave sediments; 
2 packrat 

24 000 to 4000 

High above cayon bottom, 
relatively mesic site, 
pinyon-juniper with 
scattered subalpine 
conifers, little active talus 



1640 m 

Small overhang; 
2 packrat 

13 500 to 12 000 

Small rock outcrop in the 
middle of Snake Valley, 
desertscrub, little active 

Packrat middens. — All fossil packrat middens in this report were well-indurated 
with urine. To insure that stratigraphic units of different ages were not mixed during 
sampling, each midden was examined carefully prior to collection. Approximately two 
kilograms were removed from each stratigraphic unit and packaged in the field. In the 
laboratory approximately one kilogram of each unit was soaked in water until the 
cementing urine dissolved. The disaggregated samples were then washed through 20- 
mesh screens and oven dried. The fossils were hand-sorted, and after identification, 
plant remains were selected for radiocarbon dating and pretreated with 10% HCI in an 
ultrasonic cleaner. Radiocarbon dates from the packrat middens and the cave sedi- 
ments (discussed below) are presented in Table 2. 

Four of the 5 packrat middens recovered from Smith Creek Cave will be discussed. 
Radiocarbon dating of these middens indicates a time range from 1 1 650 ± 280 to 
13 340 ± 430 yr B.P. (years before present. Table 2). Ladder Cave, a much smaller 
cave, is located immediately above Smith Creek Cave. We discuss 6 packrat middens 
from this cave, dating from 1 1 080 ± 115 to 27 280 ± 970 yr B.P. (Table 2). Two packrat 
middens from Council Hall Cave (Table 2) provided faunal remains of middle Holocene 
age (4220 ± 60 to 6120 ± 80 yr B.P.). Three middens reported from the Streamview 
locality date from the middle Holocene (6490 ± 190 yr B.P.), late glacial (1 1 010 ± 400 
yr B.P.), and the end of the last full-glacial (17 350 ± 435 yr B.P.). Amy's Shelter is 
in the bottom of the canyon across the canyon from Smith Creek Cave. This site and 
Kachina Cave are included here to add the results of Miller's (1979) faunal study. 

Smith Creek Cave stratigraphy.— The stratigraphy currently known from the 

Table 2. Radiocarbon dated packrat middens and cave sediments from Smith Creek Canyon, Snake Range, 
Nevada (Thompson 1979, Thompson and Mead 1982). Associated plant material: A) Dung and/or 
unidentified plants; B) Ephaha vindis\ C) Juniperns communis: D) J. osteosperma; E) Picea enfielmannii; 
F) Pinu.s fli-.xilis; G) P. lonf^iwva: H) P. monophvlla: I) Artemisia spp.; J) Afriplex confertifolia; K) 
Cercocarpus intricatus; L) Chamuchatiaria millijoliiim: M) Chrysothumnus sp.; N) Foresellesiu neva- 
densis; O) Ribes montif^eniim; P) Symphoricarpos sp.; Q) Rhus triolhala. 

Radiocarbon age 




Lab no. 

(yr B.P.) 



Council Hall Cave (CHC) 















Ladder Cave (LC) 



11 200 







27 280 







17 960 







13 230 







12 100 












Smith Creek Cave (SCC) 










1 1 650 







12 235 







13 340 





(Reddish-brown Silt: SCC Sed.) 


28 650 





Streamview Shelter (STV) 










17 350 












northwestern sections of Smith Creek Cave was estabHshed from the excavation of 
Test Pits 2 and 3 (Bryan \979a). The stratigraphic units established by Bryan will be 
followed in this report. Bryan recognized 3 stratigraphic "zones" in the rear of Smith 
Creek Cave, the lowest unit being the Cemented White Silt Zone. In the northwestern 
section of the cave, this unit was eroded away before the deposition of the second 
stratigraphic zone — the Reddish-brown Silt Zone. Bryan noted that the erosional dis- 
conformity between the Cemented White Silt Zone and the Reddish-brown Silt unit is 
near vertical. The reddish silt of the latter unit is believed to be of probable eolian 
deposition, although it is just as likely that water from a nearby ceiling conduit may 
have caused the erosion of the first stratigraphic unit and at least a partial deposition 
of the Reddish-brown Silt. This latter unit is very fossiliferous (partially a raptor ac- 
cumulation) and is stained reddish-brown by the surrounding silt (dry 7/6 lOYR. yellow; 
damp 5/8 7.5YR, strong brown, Munsell color; our observations). No organic remains, 
including charcoal, were observed in the Reddish-brown Silt by Bryan (\979a) or us. 
Permineralization has occurred with all of the red-stained bones (Miller 1979). Younger 
contaminating bones are white, not mineralized, and easily spotted. 

Bryan ( 1979a) reported that "a sample of unidentified red-stained bone scrap [from 
Test Pit 2] yielded a collagen date of 28,650 ± 760 years B.P. (Tx- 1639)." A discrepancy 
arises in that Valastro (1977) reported that the Tx-1639 '^C date is from a charcoal 
sample from Test Pits 2 and 5; Test Pit 5 is at the mouth of the cave. We did not find 
charcoal in the unit in Test Pit 2. Because the red-stained bones in the Reddish-brown 
Silt in Test Pit 2 is permineralized with the loss of most bulk organic constituents 
(Miller 1979), we assume that the radiocarbon date from the unit is at best a very rough 
estimate of its true age and that the material dated was probably bone scraps, not 
charcoal. Further radiocarbon dating of this unit is in order. 

Bryan indicated probable temporal correlations between the Reddish-brown Silt 
and the Laminated Pink Silt and Rubble Zone recognized in the excavation of the 
deposit at the mouth of the cave. Unlike the Reddish-brown Silt, this middle unit at 
the cave entrance rarely contains bones (Bryan 1979a). Although no radiometric dates 
were obtained from the Laminated Pink Silt and Rubble Zone, a '^C date of 12 600 ± 
170 B.P. (A-1565) was obtained at the lower boundary of the above unit — Bristlecone 

Table 3. Wisconsinan and early Holocene fish, amphibians, and reptiles from Smith Creek Canyon. G) = 
previously reported by Goodrich (1965). B) = previously reported by Brattstrom (1958, 1976). 1) = now 
living within 50 miles; 2) = not living locally but elsewhere in Great Basin; 3) = not presently within the 
Great Basin; ? = age assignment in question. 



s >> 

1/5 c 

Oi o 

















o ^^ 

oo c 
n o 

A\ -^ 













E a" 

on xi >^ 





^ o 










Salmo clarki 
Gila atraria 


Scaphiopiis cf. intermontanus X 

5. cf. hammondi (B) 3 

fiw/o boreas 2 

Bufo cf. woodhousei 1 

Bm/o sp. (B) 
/?a«a sp. 


Crotaphytus collaris X 

Crotaphytiis wislizeni X 

Crotaphytus sp. 

Sceloporus magister 2 

Sceloporus occidentalis or iindulatus X 

Sceloporus graciosus (B) X 

Sceloporus sp. 

f/Za stansburiana (B) X 

Phrynosoma platyrhinos (B) X 

Phrynosoma douglassi 2 

Phrynosoma sp. (G) 

Cnemidophorus cf. //^r/5 X 

Coluber constrictor (G, B) X 

Masticophis flagellum (B) 2 

Pituophis melanoleucus (G, B) X 

Lumpropeltis getulus (G, B) X 

Lampropeltis pyromelana X 

Lampropeltis triangulum 3 

Rhinocheilus lecontei X 

Thamnophis sp. (G) 

Hypsiglena torquata (G, B) X 

Crotalus cf. viridis (G) X 

Crotalus viridis (B) 















































Pine and Sheep Dung Zone (Bryan 197%/). Presumably this radiocarbon date provides 
a minimum age estimate for the Laminated Pink SiU and Rubble Zone and by corre- 
lation the Reddish-brown Silt unit. 

The third depositional unit Bryan described from the rear of the cave is the Grey 
Silt, Rubble, and Dung Zone. Organic remains are very common within this unit. All 
bones incorporated in this uppermost unit are various shades of white without per- 
mineralization. Occasional mixing of the red-stained bones by bioturbation has oc- 
curred near the lower boundary of the unit. Although this unit has not been radiocarbon 

dated, it certainly is younger than 10 000 yr B.P. and the upper portion probably is 
less than 3000 years old (Bryan 1979^/). 


Table 3 is a chronological check-list of late Pleistocene and Holocene age fish, 
amphibians, and reptiles from Smith Creek Canyon. Many of these taxa have not been 
reported previously from the late Pleistocene of the Great Basin. Table 4 is an updated 
check-list of fossil mammals known from Smith Creek Canyon. Although previous 
reports have described the mammals (Harrington 1934. Stock 1936. Goodrich 1965. 
Miller 1979). this table shows their stratigraphic provenience, not adequately given 
before . 

Following is an annotated list of the fossil fish, amphibians, reptiles, and mammals 
that we recovered from packrat middens and from one sedimentary layer (Reddish- 
brown Silt) in Smith Creek Cave. Abbreviations for the fossil localities are listed in 
Table 2. The number in parentheses refers to the quantity of that element. We follow 
the nomenclature and ordering, unless otherwise stated, of Smith (1978) for the fish. 
of Stebbins (1966) for the amphibians and reptiles, and of Jones et al. (1979) for the 

Class Osteichthyes 

Family Salmonidae 

Salmo clarki (Cut-throat Trout) 

Material. — SCC Sed.: R angular. R dentary. vertebrae (25). 

Remarks. — The right angular resembles that of a small individual of the Great 
Basin, not of the Colorado River. The right dentary is of a specimen of approximately 
100 mm in length. All the vertebrae are of small individuals. The skeletal elements 
were identified as Salmo rather than Salvelinus malma (Dolly Varden. the other trout 
in the Great Basin) because ( 1) the angle between the coronoid and post-dorsal process 
of the angular is near 90°, and (2) the dentary lacks the deep groove under the tooth 
platform. The small size of the specimens suggest that they were from a creek (Smith 
Creek?), not a lake (Lake Bonneville) (Gerald R. Smith identifications and personal 
communication 1981). 

Distribution. — Salmo clarki, which occurred in Lake Bonneville during the Pleis- 
tocene, is the widespread trout of the Great Basin and Intermountain Region (Smith 
et al. 1968. Smith 1978). 

Family Cyprinidae 
Gila atraria (Utah Chub) 

Material. —SCC Sed.: Basioccipital and vertebrae (3). 

Remarks.— The basioccipital with a pharyngeal process has the shape and angles 
of Gila atraria, not Richardsonius helteatiis (Redside Shiner), which has a less ovoid 
haemal canal and a less obtuse angle between the cranial part o\' the bone and its 
pharnygeal process. The basioccipital belonged to a fish about 1 10 mm long (Gerald R. 
Smith identification and personal communication 1981). 

Distribution.— Gila atraria is native to the Bonneville and upper Snake River 
drainages and occurred in Lake Bonneville during the Pleistocene (Smith et al. 1968. 
Smith 1978). 

Class Amphibia 

Order Salientia 

Family Pelobatidae 

Scaphiopus cf. intermontanus (Great Basin Spadefoot Toad) 

Material.—SCC 1: tibiofibula; SCC Sed.: tibiofibulae (13). radio-ulnae (3). 
Remarks. — No comparative material was available for Scaphiopus intermontanus 
although we have an excellent series of the closely related S. luimmondi. 

Distribution. — Only this species of spadefoot toad presently occurs within wood- 
lands and sagebrush areas in the Great Basin (as well as in the Snake Range); however, 
S. cf. hammondi has been identified from an unprovenienced level in Smith Creek 
Cave (Brattstrom 1976). ^ 

Family Bufonidae 
Biifo bore as (Western Toad) 

Material. — SCC Sed.: sacral vertebra, R ilium. 

Distribution. — Bufo horeas presently occurs throughout most of the Great Basin 
except the drier eastern part along the Nevada-Utah border; it has not been found in 
the Snake Range. 

Bufo cf. woodhousei (Woodhouse's Toad) 

Material.— SCC Sed.: tibiofibula. 

Remarks. — Martin (1973) used the ratio of the tibiofibula, minimum width relative 
to length, to differentiate species of Bufo {B. boreas: 690-760; B. cognatus: 840-1050; 
B. hemiophrys: 950-1160; B. microscaphus: 670-850; and B. woodhousei: 770-950). 
The ratio of the Smith Creek Cave sediment fossil is 1013 (1.84/18.16 mm x 10 000), 
being relatively stout and thick in the middle. Although our fossil could be identified 
as B. cognatus based upon the tibiofibula ratio, we refer our specimen to B. wood- 
housei because of its present closer distribution; B. cognatus does not live in the Great 
Basin. Additional fossils of Bufo are needed to refine our identification. 

Distribution. — Bufo woodhousei presently does not inhabit much of the Great 
Basin; the western edge of its range is just north and east of the Snake Range. 

Bufo sp. (toad) 

Material. — SCC Sed.: tibiofibulae (3), radio-ulna. 

Remarks. — We were unable to identify the fragmented fossils to species. 

Family Ranidae 
Rana sp. (frog) 

Material. — SCC Sed.: tibiofibulae (4), coracoid, atlas. 
Remarks. — We were unable to assign the specimens to species. 
Distribution. — Presently R. pretiosa and R. pipiens occur within the Great Basin 
(Stebbins 1966). 

Class Reptilia 

Order Squamata 

Suborder Sauria 

Family Iguanidae 

Crotaphytus collaris (Collared Lizard) 

Material. — SCC Sed.: L dentaries (3), R dentary. 

Remarks. — C. collaris and C. wislizeni can be separated from most other iguanid 
lizards by their larger size and their dental characters. The individual teeth of C. 
collaris are relatively wide anteroposteriorly as compared to those of C. wislizeni, the 
posterior teeth strongly tricuspid, the anterior tending toward blunt spikes, some with 
a slight posterior curve. In C. wislizeni the anterior three-quarters of the teeth are 
sharp, recurved simple cusps with only a few posterior teeth tricusped. 

Distribution. — The Collared Lizard occurs throughout the Great Basin in a variety 
of mountain and rocky habitats. 

Crotaphytus wislizeni (Leopard Lizard) 

Material. — SCC Sed.: L dentaries (2), L maxillae (3). R dentaries (3), R maxillae 




Figure 2. Present distributions of Sceloporus magister (L's), Lampropehis triangiiliim (parallel lines), 
and L. pyroineUina (mixed lines) (from Stebbins 1966). ( I) Fossil localities and Snake Range. The distribution 
of L. triangulum overlaps the entire range of L. pyromelana only in Utah. The distribution of .V. magister 
overlaps only the southern tip of both species of Lamprupeltis in Utah and all of L. pyronwlanti in Arizona. 

Remarks. — The distinguishing characters are listed under C. collaris. 

Distribution. — Like the Collared Lizard, the Leopard Lizard occurs throughout 
the Great Basin, although usually in desertscrub communities on the finer alluvial 
habitats of the valleys. 

Crotuphytus sp. (Collared or Leopard Lizard) 

Material. — SCC Sed.: L dentaries (2), L maxillae (3), R dentaries (2), R maxilla. 
Remarks. — We were unable to assign these elements to species. 

Sceloporus magister (Desert Spiny Lizard) 

Material. —SCC Sed.: L dentary. 

Remarks. — The Desert Spiny Lizard can be differentiated from other sceloporine 
lizards of the Great Basin by its larger size. 

Distribution. — The range of S. magister presently ends southwest of the Snake 
Range (Fig. 2). Although no other spiny lizards reach the size of 5. magister in the 
Great Basin, other very similar sized species occur farther south in southern Arizona 
(S. clarki and S. Jarrovi). 


Sceloporus occidentalis or undiilcitiis (Western or Eastern Fence Lizard) 

Material. — LC 2a: epidermal scale; SCC Sed.: L dentaries (6), R dentaries (2), R 
maxillae (5), frontal. 

Remarks. — There are no satisfactory dental criteria for separating the closely re- 
lated S. unditlatiis and S. occidentalis; both species have larger races at their northern 
distribution. Since the latter species now lives within the Snake Range, we refer our 
fossils to it. The nearest population oi S. undidatus is to the east in Utah. Both species 
possibly occurred within the mountain range in the late Pleistocene. 

Sceloporus graciosus (Sagebrush Lizard) 

Material.— SCC Sed.: R dentaries (2), frontal. 

Remarks. — The skeletal elements of 5. graciosus can be differentiated from those 
of other sceloporine lizards of the Great Basin by their distinctly smaller adult size and 
their slender pointed teeth with weakly developed secondary cusps. 

Distribution. — S. graciosus now occurs throughout the Great Basin including the 
Snake Range. 

Sceloporus sp. (spiny lizard) 

Material. — SCC Sed.: L dentaries (2), L maxilla. R dentary, R maxillae (2). 
Remarks. — We were unable to assign these elements to species. 

Uta stansburiana (Sideblotched Lizard) 

Material. — LC modern: L dentary. 

Remarks. — The single element compared well with modern Uta stansburiana. The 
lizard presently lives in Smith Creek Canyon, and the only skeletal specimen recovered 
in our study came from a modern packrat midden. 

Phrynosoma douglassi (Short-horned Lizard) 

Material.— hC 2a: frontal; LC 3: L maxilla; SCC Sed.: L dentary, R maxillae (2), 

Remarks. — The recovered dentary, maxillae, and frontals could not be distin- 
guished from those of modern P. douglassi. In bone and dental characters of the 
frontal, dentary. and maxilla, P. douglassi are distinct from P. platyrhinos (Robinson 
and Van Devender 1973). P. douglassii has tall Sceloporus-WkQ teeth with well-devel- 
oped secondary cusps and a rounded bottom to the dentary. P. platyrhinos has short 
peg-like teeth with reduced secondary cusps. 

Phrynosoma platyrhinos (Desert Horned Lizard) 

Mrt/mV//.— SCC Sed.: L dentary. 

Distribution. — Both Phrynosoma platyrhinos and P. douglassi live within the 
Great Basin today, the latter only at higher elevations on a few mountain ranges in 
northeastern Nevada (Stebbins 1966). 

Family Teiidae 
Cnemidophorus cf. //^t,'/7.v (Western Whiptail Lizard) 

Material. —SCC Sed.: R dentaries (4). 

Remarks. — We refer our specimens to C. cf. tigris because it is the only species 
of whiptail lizard now living within the interior Great Basin. Modern C. tigris and the 
fossils have very large dentaries. Other species of whiptail (e.g., C. hurti) with den- 
taries of similar size now occur far south of the Great Basin. 

Suborder Serpentes 

Family Cokibridae 

Pituophis fucldnolenciis {Gt>pher Snake) 

Material. see Sed.: palatine, vertebrae (23). 

Ri'inarks. — The vertebrae o'i Fituoplus inelanolcucus are most similar to those of 
Elaphe. We used the criteria described by Auffenberg (1963) to differentiate the two 

Distribution. — The Gopher Snake is very common throughout the Great Basin. 

Lampropeltis pyromeliimi (Sonoran Mountain Kingsnake) 

Material.— sec Sed.: vertebrae (29). 

Remarks. — The vertebrae of Lampropeltis pyromelana are typical of kingsnakes, 
with well-developed subcentral ridges. We differentiate L. pyromelana from L. trian- 
gidum by two criteria: (1) the cotyle and condyle are proportionally larger in L. py- 
romelana at all stages of growth, and (2) the accessory processes are fairly pointed on 
L. pyromelana whereas they are longer and globose (as viewed anteriorly) on L. trian- 
gulum. Both species have low neural spines unlike L. getuUis or Rhinoeheilns leeontei. 

Distribution. — The Sonoran Mountain Kingsnake presently occurs only in a few 
relictual populations in the Great Basin including the Snake Range (Fig. 2). It com- 
monly occurs in the central and southern Wasatch Mountains and the mountainous 
region farther south and west. 

Lampropeltis triangulum (Milksnake) 

Material. — SCC Sed.: vertebra. 

Remarks. — The distinguishing criteria were discussed under L. pyromelana. 

Distribution. — The Milksnake occurs on the eastern periphery of the Great Basin 
in the Wasatch Mountains and farther south in Utah but not as far west as the Snake 
Range (Fig. 2). 

Rhinoeheilus leeontei (Long-nosed Snake) 

Material. — LC modern: vertebra; SCC Sed.: vertebrae (13) 

Remarks. — The vertebrae of Rhinoeheilus are quite characteristic and can readily 
be identified (Hill 1971, Van Devender and Mead 1978). 

Distribution.— The Long-nosed Snake lives throughout the Great Basin except for 
the northwesternmost region. 

Thamnophis sp. (garter snake) 

Material.— SCC Sed.: vertebrae (3). 

Remarks. — We were unable to assign the specimens to species. 

Distribution.— Presently T. elegans (Wandering Garter Snake) lives within the 
Great Basin, including the Snake Range, and T. sirtalis (Common Garter Snake) bor- 
ders parts of the basin (Stebbins 1966). 

Hypsiglena torquata (Night Snake) 

Material.— CHC lb: vertebra; SCC Sed.: L dentary. vertebrae (36). 

Remarks.— H. torquata has small generalized colubrid vertebrae, which are very 
similar to those of other small snakes such as Sonora semianulata. Identification cri- 
teria used here were those used by Van Devender and Mead (1978). 

Distribution. — The Night Snake occurs over most of the Great Basin. 




(6/.6I Aqonj. 

1) T3 ™ 




•6Z.6I J3n!P\) dO J-^ 

U II 'o 




(X)i:-009 (<•.) sjBiuixojddv 

■S^ c 


(6Z.6I UMiJO 



•6Z.6I J3|I!W) d'O J-^ 




(<;) 3iBiuixojddv 




(q6Z.6l UB'^JO 

• E :2 





'6Z.6I JSiniAl) USAJS 

(U « w 




)0u aSy  

da j^oooiT^*. 




. (*) = 
stion t 



jaqjo puB 

SUHQ puB 3 

SllUn 3U3D010H 

da J-^ 000 013= 
qqna jus -^sjo 



is te 
rs q 



(1X31 33S) do -"^ 000 cl« 
ins uMOjq-qsippay 

o ji: w 


^ '^ •£ 



D. C o 



S[3A3| [JB p3ZI|EJ3U39 

C3 ■- 
u — , u 



yodsj sinx 

i: 3 i 


he s 
We ; 

6Z.6I Jail'H 

H -u 

• — 

. ^ II 


ents ; 
. (?) 


596 1 qoupooQ 


000 8c-000 LZ 

- M ^ 

c •— c 

1 ^c1 

000 81-000 Z.1 

§ w -^ 


^ ^U 



1 Crt 
): th 







■t: ^ c/3 



.t: ON 





000 21-000 11 

1 =^ 




2 <« S 




000 11-000 01 

c/3 OQ Cd 




d by 




2 <u t^ 



E rS O 



w b 2? 


c 5; c 


0) oj 'z: 



— K a. 


o u. J7 


K 2 ^ 



31IBA 31Eip9lUUJI 

JO aSuB^ 3)jEUS ui Ajjuasajj 

§ 5== 



■- ^ ^ 





-^ J 2i 



^<2 '- 


^ f 


UJ ^ O 


-1 ^ D. 


«u 2i 





^\ <S ^ ?S ^ ?S 

^- X X X X 


X ^- X x^-x 

X X 



X X 

X X 




X X 




X X 







"^ Ci. ii 





i: 3; 5: t*- 

•S c • 

iO C to 

"1. S 


'-> S 



s: — 

o ^ 


































c« CD 

>> c 

c ■"" 


JZ T3 





(6/.61 ^qonj. 

■6Z.6I J3|l!l\) da J-^ 

00i:-009 (<,)3>BUJ!XOjddv 

(6Z.6I uqiJO 

'6Z.6I J3|l!l\) da J-^ 

OOOS-OOSI (<.) aiEuiixojddv 

(q6/i6i u'^'^Ja 

lou aSv "da ■''< 000 iZ^i 

SJlUn 3U3DO10H 

jamo pire da J-^ 00001 s 
3una put? 3|qqny -JUs AajQ 

(IX3J 33S) de J-^ 000cl=£ 
l|iS uMOjq-qsippay 

S|3A9I n^ — P3ZI1BJ3U39 

uodaj Sim 

6/.6I JainW 
S%I qaupooo 
000 gc-OOO At 
000 81-000 Z.I 
OOOcl-000 11 


000 01-0008 


XaijBA ajBipsuiLU! 

JO 33UB^ 351BUS Ul A[JU3S3Jd 


X X 



X X 


X X 



xxxxxx xxxxxxxx xxxxx 



X X 


xxxxxx X XX 



X X 




X X X ^-X X X 

X X 

X X 
X X 


X X X X X X X 

X X 

X X 



D. 5 


^ s 


'-> >1 z 


>^ -^ :: z 

^ ^ -• ^ s ^ 

^ -^ ^. »*. - 

ci. •-- S S i^ 2 

-Cj n3 '^ '^ i^. i 

' ^ i; 5 S ~ "J 

■i: t Ci.A i' 

w^ .?" ,^ >* 

~. ^ -* 


•^ ^— "^ c - 

S 5 '^ S t 

r* *-• w ^ *» 

§ qqq::?:^^^^: 




















































(J M 

(6Z.6I ■<Moni 

■6Z.6I J3|I!l\) da -"^ 

(X)i:-009 ((,) aiRiuixojddv 

(6/.61 "MiJO 

'6Z.6I J3|l!l\) da J-^ 

OOOs-OOsI (('.) aiBiuixojddy 

(q6/.6i uR^Ja 

'6Z.6I J3|!!IM) "3AiS 

jou asy da J'^ oooe:=£<. 

SJIUn 3U33010H 

jaqjopuB da J^ 000 013= 
3una puB aiqqny 'ins XajQ 

(1X3J 33S) je J-^ 000cl=£ 
ins UAVojq-qsippay 

S[3A31 [[B — p3Zl|BJ9U3r) 

jjodaj siqx 

6/.6I J^II'PM 

000 8c-000 Li 
000 81-000 Z.I 


OOOcl-000 11 

000 1 1 -000 01 



X3[|BA ajBipaiuiui 

JO aSUBy 3}(BUS UI /(|}U3S3J(J 





X X 


x^-xx^-xxxx xxxxxxxxx^-x 


X X 



X XXX X x^- x^-xx 



X ^-x 




a ^^ 




5 * 





Canis cf. latra 
C. cf. lupus 
Vulpes vulpes 
V. velox 
Ursus sp. 

Bassariscus as 
Martes nobilis 
Martes sp. 
Mustela ermin 
M. frenata 
M. vison 
Mustela sp. 
Taxidea taxus 
Spilogale puto 
Spilogale sp. 
Felis rufus 
F. cf. onca 
F. onca 
F. concolor 
IF. atrox (*) 
Equus sp. (Lai 



«— ' 




as u 


C > 



[^ <U 

>^ — 

E w 


o ffi u 




-^ CU 

c ■" 
U op 





(6/.6I '^Moni 

•6/.6I J3|l!l\» de -"^ 


(6A6I uqnji) 

•6Z.61 J3|I!l\) dO J< 

IK)0!;-()0!,I (i.) ajEUJixojddv 

(q6Z.6l u'^'^JO 

■6Z.61 J3|I!IA1) ua-^lS 

jou aSv da J-^OOOEc^c 

sjiun auanojOH 

jsqjo puB de JA oo()Ols 

Suna pup 3|qqny -ills '^SJO 

(JX31 33S) da ja 000 :i« 

JlfS u/v\ojq-qsipp3y 

S|3A3| IIB P3ZI1BJ3U39 

jjodaj sm± 

6Z.6I J3II!IM 

^"96 1 M^upooo 

000 8:-000 LZ 

000 81-000 AI 


000 £1-000 :i 


000 11-000 01 

000 01-0008 


0009-000 1' 

A3||EA aiRipauiuji 
JO aSucy 35(bus u; Xijuasajj 


X X 


X X 

xxxx^-x ^-XXX X X 


X X 






<r- 55 

^ p. 


5^ = - 

y. i ^ -^ -^ 

5 S S r ^ ^ "t •'^ :- S. ? ? 'S .2 'B -2 
Uj (o c"- O O 'j; 'J < ;■• c- O ^ o O c^- O 


Family Viperidae 
Crotalus cf. viridis (Western Rattlesnake) 

Material. see Sed.: vertebrae (5) 

Distribution. — We refer our specimens to C. viridis because they are from a me- 
dium-sized rattlesnake and this species presently inhabits the Great Basin. C. mitchelli 
is very similar but occurs no farther north than the Mohave Desert. 

Class Mammalia 

Order Chiroptera 

Genus and species indeterminate 

Material. — SCC Sed.: isolated teeth, mandible. 

Remarks. — The isolated teeth and the fragment of a mandible did not allow generic 

Distribution. — Many species of bats occur in the Great Basin (Barbour and Davis 

Order Lagomorpha 

Family Ochotonidae 

Ochotona cf. princeps (pika) 

Material.— SCC 4: dung pellets (17), RM>; SCC 5: dung pellets (2), RP; SCC 
Sed.: LP3; STV 1: dung pellet; STV 2: dung pellets (>50); STV 3: dung pellets (3). 

Remarks. — The RP^ from SCC 5 is referred to Ochotona, but in view of its worn 
state, it could possibly be a Sylvilagus idahoensis. Miller (1979) reported that pika 
remains were recovered in all stratigraphic units in Smith Creek Cave; the youngest 
occurrence, however, cannot be determined. Pika also was recovered from Council 
Hall Cave; unfortunately, the remains (not described) were not reported in relation to 
the testpits, stratigraphy, or associated radiocarbon age (Miller 1979, Bryan \919b). 
Small mammal bones {Ochotonal) occurred throughout the upper two meters (above 
a 23 900 ± 970 yr B.P. radiocarbon date, GaK-5100) of organic layers in Test Pit 2 
(Bryan 1979/?). 

Two packrat middens in Smith Creek Cave and three middens in Streamview Shelter 
contained lagomorph dung pellets referable to Ochotona cf. princeps (Fig. 3). Dung 
pellets from each packrat midden were measured (length and width) and the measure- 
ments compared with those from samples from modern Ochotona princeps, Sylvilagus 
idahoensis, and S. nuttallii. The advantage of the preserved dung pellets from the 
packrat middens over the skeletal fragments from the cave sediments is that presum- 
ably Ochotona lived at the fossil site and was not brought there via a raptor stomach. 
Pika (skeletal remains) has been identified previously from mountain ranges outside its 
present distribution and from elevations below its current lower limit in ranges it now 
inhabits (Fig. 4; Grayson 1977, 1981, in press a, b. Miller 1979). To our knowledge, this is 
the first record of late Pleistocene age dung of Ochotona south of the permafrost in 
North America (Guthrie 1973). A more detailed account of the fossil Ochotona in the 
Great Basin is in progress. 

Distribution. — Pika does not now live in the Snake Range or in any nearby moun- 
tains (Hall 1946, Hall and Kelson 1959), although it does inhabit the mountainous 
regions on the west, north, and east sides of the Great Basin (Fig. 4). Only O. p. 
nevadensis (Ruby Mountains region) and O. p. tutelata (Toquima Range and Desatoya 
Mountains) live in restricted relictual localities within the interior Great Basin. 

Family Leporidae 
Sylvilagus {^Brachylagus) idahoensis (Pygmy Rabbit) 

Material.— SCC Sed.: M,,. 

Remarks. — S. idahoensis has not been previously recorded from Smith Creek 
Canyon (Miller 1979). In the central Great Basin at Gatecliff Shelter, the Pygmy Rabbit 






m m 

Figure 3. Measurements (width and length) of lagomorph dung pellets, (a) Ochotona cf. princeps. fossil, 
this report; (b) O. princeps, modern, California; (c) Sylvilugus idahoensis, modern, Nevada: (d) S. nuttallii, 
modern, Nevada. 

was recovered throughout the deposit (Grayson /// press h). North of the Great Basin, 
S. idahoensis is recorded in late Pleistocene context at Jaguar Cave (Guilday and 
Adams 1967), Wasden site (Owl Cave, Guilday 1969), and Moonshiner Cave (Anderson 
1974). South of the Great Basin it is reported from Tule Springs, Nevada (Mawby 1967) 
and Isleta Cave, New Mexico (Harris 1977). 

Distribution. — The Pygmy Rabbit presently occurs across the northern and east- 
central Great Basin (Hall 1946). 

Sylvilagus sp. (rabbit) 

Material. —see Sed.: RP^ 

Remarks. — The specimen could not be identified to species. 

Lepiis sp. (hare) 

Material.— sec Sed.: RM,. 

Remarks. — The specimen could not be identified to species. 


Figure 4. Distribution of Ochotona princeps on isolated mountains within and bordering the Great Basin 
(from Hail 1946, Hall and Kelson 1959). (1) Fossil localities of Smith Creek Canyon and Garrison. 

Order Rodentia 

Family Sciuridae 

Eiitamias minimus (Least Chipmunk) 

Material.— sec Sed.: R mandible P4Mi_3. 

Remarks. — This is the smallest chipmunk in Nevada. Assignment of our specimens 
to this species was on the basis of size. Although the other species of Eutamias of the 
Great Basin are restricted to the vicinity of coniferous trees, E. minimus may live in 
sagebrush {Artemisia tridentata) at both high and low elevations (Hall 1946). 

Eutamias cf. umhrinus (Uinta Chipmunk) 

Material.— sec Sed.: frontal. 

Remarks. — Neither of the two species of Eutamias have been reported from Smith 
Creek Canyon (Miller 1979). Our specimen compared most favorably with modern E. 

Distribution. — The Uinta Chipmunk is a medium-sized chipmunk that presently 
occurs in the Snake Range (Hall 1946). 


EutiiDiias sp. (chipmunk) 
Material. see Sed.: LM, (2). RM,. RM' (3), LM-', RM^ 

Mannota flavivcntris (Yellow-bellied Marmot) 

Material. see Sed.: LI. 

Remarks. — Although we are assuming our specimen is M.flaviventris, the isolated 
incisor does not allow for specific identification. The Hoary Marmot, Mannota cali- 
gata, cannot be ruled out definitively based upon our specimen, though its occurrence 
in the Great Basin in the late Pleistocene seems unlikely. 

Distribution. — Marmota jlaviventris presently occurs in the Snake Range at ele- 
vations higher than Smith Creek Cave. 

Ammospermophilus lencnrns (White-tailed Antelope Squirrel) 

Material.— Le 2a: frontal; SCC Sed.: L mandible M,_.^,. 

Remarks. — The White-tailed Antelope Squirrel has not previously been reported 
from Smith Creek Canyon (Miller 1979). Our specimens compare with modern repre- 
sentatives of y4. lencnrns. 

Distribution. — Ammospermophilns lencnrns presently lives in most of the Great 
Basin, including the area of the Snake Range (Hall 1946). 

Spermophilns cf. richardsonii (Richardson's Ground Squirrel) 

Material.— 'Le 1: RM^; LC 4: LM,. 

Remarks. — Spermophilns richardsonii has not been recovered previously from 
Smith Creek Canyon, though it has been recorded from the more northern localities 
of Jaguar Cave (Guilday and Adams 1967) and Moonshiner Cave (Anderson 1974), 
Idaho. Both Davis (1939) and Hall (1946) have expressed the belief that S. richardsonii 
must have occupied a much wider range over an ecological area now filled by S. 
beldingi and S. armatns. On the basis of the Smith Creek Canyon fossils of S. cf. 
richardsonii, we concur with them. Our two specimens compared most favorably v\ith 
modern S. richardsonii. 

Distrihntion. — Richardson's Ground Squirrel does not occur presently in the 
Snake Range. The present distribution of 5'. r. nevadensis in the Great Basin is centered 
in the Independence Mountains of northern Nevada, and it occurs no farther south 
than the Roberts Mountains (Hall 1946). 

Spermophilns cf. beldingi (Belding's Ground Squirrel) 

Material.— Le 2b: L maxilla M'. 

Remarks. — Spermophilns beldingi has not been reported previously from Smith 
Creek Canyon, and the only other published fossil or subfossil record for Belding"s 
Ground Squirrel in the Great Basin is from Stratum 3 at Gatecliff Shelter (Grayson //; 
press h). Our single specimen compared most favorably with modern ,S. beldingi. 

Distribution. — Spermophilns beldingi is ecologically and physically very similar 
to S. richardsonii. Belding's Ground Squirrel presently does not occur in the Snake 
Range or in any immediate mountain range. Its present distribution centers in the 
northern Sierra Nevada and in the higher mountains of north central and northern 
Great Basin, though it occurs farther south than .S'. richardsonii. dow n to the TcHiuima 
Range (Hall 1946)'] 

Spermophilns cf. lateralis (Golden-mantled Ground Squirrel) 

Material. — LC modern: RM,; LC 2a: L maxilla M'"''. L mandible M,.;,; SCC 1: 
R mandible P,M,; SCC Sed.: R mandible M,_,. RM,. LM'. RM,_.,. 

Remarks. — Although S. lateralis was not previously reported from Smith Creek 
Canyon (Miller 1979). it is fairly widespread in the Smith Creek Canyon fossil localities 


and apparently was common in the canyon throughout most of the late Pleistocene. 
Although we have found this to be the most common species of Spermophiliis, Miller 
(1979) considered that S. cf. tcnvnsendii was the most prevalent in the Smith Creek 
Cave record. Miller (1979) stated that Goodrich (1965) referred his specimens from 
Smith Creek Cave to S. cf. townsendii; although he considered that species, he did 
not definitely refer his specimens there. However, the presence of Townsend's Ground 
Squirrel in Smith Creek Canyon in the late Pleistocene seems plausible. 

Spermophilus sp. (ground squirrel) 
Material.— \.C 2b: RM,; SCC Sed.: P (3), RMg, RM^. 

Family Geomyidae 
Thomomys sp. (pocket gopher) 

Material. —SCC Sed.: LM». 

Remarks. — We were unable to assign our single specimen to a species. 

Family Heteromyidae 
Perognathus sp. (pocket mouse) 

Material. —SCC 1: L mandible P4 (3), R mandible, R maxilla M^; SCC Sed.: L 
mandible, R maxilla M* (2); STV 2: L maxilla (2), R maxilla. 

Remarks. — We were unable to assign these specimens to species. 

Dipodomys sp. (kangaroo rat) 

Material.— SCC Sed.: LP. 

Remarks. — We were unable to assign this specimen to species. 

Family Cricetidae 
cf. Peromysciis (white-footed mouse) 

Material.— CnC lb: RM'-^; SCC I: LM^; SCC Sed.: LM', RM'-^; STV 2: LM,, 
RM'-^: STV 3: LM,_2. 

Remarks. — We have not identified these specimens to species because we do not 
have available a sufficient comparative collection of Reithrodontomys and Peromyscus 
from Nevada. Presently Reithrodontomys megalotis, Peromyscus eremicus, P. crini- 
tus, P. maniculatus, P. hoy Hi, and P. truei live in or near the Great Basin. Previous 
work at Smith Creek Cave has produced specimens assigned to Peromyscus sp., but 
Reithrodontomys and Onychomys have not been identified from the fossil localities in 
Smith Creek Canyon, though they very well may be included. 

Neotoma lepida (Desert Packrat) 

Material.— CHC la: LM' ^ CHC lb: LM, (2); SCC Sed.: LM, (2), RM, (2), RM' 
(3), LM' (8); STV 2: RM^. 

Neotoma cinerea (Bushy-tailed Packrat) 

Material.— CHC la: LM', RM' (2); CHC lb: LM,, LM'; SCC 1: lower leg assem- 
blage with hide; SCC Sed.: RM,, RM' (7). 

Remarks. — Neotoma lepida and N. cinerea are found living in Smith Creek Can- 
yon. Their middens, both modern and fossil, can be found throughout the Great Basin. 
The excellent preservation and the advantages of having a packrat midden are illus- 
trated by the partial mummy of the hind quarters of the Bushy-tailed Packrat (ca. 
1 1 600 yr B.P.). Molars of N. cinerea are generally larger than those of N. lepida. 




Figure 5. Occlusial (left) and lingual (right) views of the Phenacoinys cf. intermedins RM, from Smith 
Creek Cave. Bar = 2 mm. 

Neotoma lepida or cinerea (Desert or Bushy-tailed Packrat) 

Material.— CHC la: RM^; CHC lb: L&RM^ (5). LM., (2). M^ (2). M, (2); SCC 
Sed.: LM, (2), RM, (2), L&RM2 (17), L&RM^ (8), M-^ (8), M3 (8). 

Neotoma sp. (packrat) 
Material.— SCC Sed.: M3; STV 2: M fragment. 

Phenacomys cf. intermedius (Heather Vole) 

Material.— SCC Sed.: RM,. 

Remarks. — Only one specimen was identifiable as Phenacomys cf. intermedius. 
Criteria for identification were the presence of tooth roots, lack of cementum, and 
distinctive occlusal pattern (Fig. 5; Guilday and Parmalee 1972). The isolated molar 
does not allow for specific identification. 

Distribution. — The Heather Vole presently does not live in the Great Basin (Hall 
1946). Our specimen from Smith Creek Cave is the first reported late Pleistocene 
Phenacomys from the Great Basin. Grayson (1981) has recovered a specimen from 
Gatecliff Shelter dating approximately 5300 yr B.P., which illustrates the middle Holo- 
cene survival of this presently extirpated mammal. 

Microtus cf. longicaudus (Long-tailed Vole) 

Material.— SCC Sed.: RM, (7), LM, (8), R mandible: STV 2: LM,. 

Remarks. — Our fossil specimens appear most similar to the Long-tailed Vole, 
although we find it difficult to differentiate M. longicaudus from M. montanus. Of the 
15 fossil M,'s examined, 14 had five closed alternating triangles, 1 had six triangles. 
Only M. montanus has been reported from Smith Creek Cave (Goodrich 1965, Miller 
1979). Goodrich compared his fossils with M. californicus, M. montanus, M. town- 
sendii, M. oregoni, and M. pennsylvanicus: M. longicaudus was not compared. 

Distribution. — Microtus montanus and M. longicaudus are the only species of 
meadow vole in Nevada (Hall 1946). Normally, M. montanus is found in the valleys 
and M. longicaudus in the mountains. 

Microtus sp. (meadow vole) 

Material.— \.C 2b: RM,_.,; SCC 4: RM,; SCC Sed.: LM' (3), RM' (3), LM^ (4), 
LM2 (4), RM3 (2). LM3 (2), RW. 


Order Carnivora 

Family Mustelidae 

Spilogale piitoiius (Spotted Skunk) 

Material.—SCC Sed.: L mandible C,P..^4M,_2. 

Remarks. — This is the only place where we disagree with the classification of 
Jones et al. (1979). We follow Kurten and Anderson (1980) and group the Western 
Spotted Skunk (Spilogale gracilis) with the Eastern Spotted Skunk {S. putorius). Our 
specimen from Smith Creek Cave has an alveolar length (anterior edge of C, to pos- 
terior edge of Mo) of 19.0 mm and an occlusial length of the Mo of 2.1 mm. These 
measurements compare well with two modern specimens of 5. putorius from the Ruby 
Mountains, Elko and White Pine Counties, Nevada. 

Distribution. — The Spotted Skunk is common throughout the Great Basin. 

Order Artiodactyla 
Genus and species indeterminate 

Material. — CHC la: dung pellets ( 15); SCC 1: keratinous hoof fragment; SCC Sed.: 
I fragment. 

Remarks. — Because of fragmented state of preservation, we were unable to iden- 
tify further these specimens. 

Discussion and Summary 

We report here the occurrence of 2 fish, 4 anurans, 9 lizards, 8 snakes, and 17 
small mammals. This assemblage adds 15 amphibians and reptiles and 7 mammals to 
the approximately 46 animals previously known from the late Pleistocene and early 
Holocene of Smith Creek Canyon (Goodrich 1965, Miller 1979). Over half of the large 
herbivores {Equus spp., Camelops sp., IHemiauchenia sp., Oreamnos harringtoni, 
and IBreameryx sp.) and the two carnivores {Martes nobilis and IFelis atrox) reported 
from Smith Creek Canyon are extinct (Miller 1979). Our data add no large herbivore 
or carnivore species to the local fauna. Of the extant species from the fossil sites. 16 
are not recorded from the Snake Range or in the immediate valleys (Bufo boreas, 
Sceloporus magister, Phrynosoma douglassi, Lampropeltis triangulum, Masticophis 
flagellum, Ochotona princeps, Spermophilus cf. richardsonii, S. cf. beldingi, Phena- 
comys cf. intermedius, Vulpes velox, Ursus sp., Mustela vison, Martes sp., Felis onca, 
Cervus elaphus, and IBison sp.). 

Few large herbivores presently occur in the Snake Range. Oris canadensis was 
reintroduced in the middle 1900s. The historic distributions of Cervus elaphus and 
Bison bison in the eastern Great Basin are not well known, though Cervus has been 
sighted in White Pine County (Hall 1946). The late Pleistocene occurrences of Ca- 
melops sp., IHemiauchenia sp., Cervus elaphus, Odocoileus sp., Equus (both large 
and small species), ?Brcameryx sp., Antilocapra americana, Oreamnos harringtoni, 
Ovis canadensis, and 'IBison in Smith Creek Canyon are fairly well documented, al- 
though the exact timing (late or middle Wisconsinan) of these occurrences is not under- 
stood and is discussed below. We question the tentative identification of Oreamnos 
americanus reported from Council Hall Cave (Miller 1979). Although the estimated 
age is not reported for the modern mountain goat specimen (it is not known historically 
from the Great Basin), we speculate that the record, if Oreamnos, is of the extinct 
Oreamnos harringtoni. 

If our reconstruction of the Smith Creek Canyon fauna-flora assemblage is correct, 
during the late Pleistocene, montane glaciers on Mount Moriah (3673 m elevation) 
moved down into the upper reaches of Smith Creek Canyon to an estimated elevation 
of 2900 m (Drews 1958). This was also a time when an open forest with Pinus longaeva, 
P. fiexilis, and Picea engehnannii was present on the coarse talus and rock outcrops 
of the limestone entrance region of the canyon. Alluvial substrates in the canyon 
bottom probably supported a mosaic o{ Artemisia spp., shrubs, meadows, and riparian 


Table 5. Fauna from two packrat middens on a rock outcrop in the middle of Snake Valley (1640 m 
elevation), near Garrison. Utah. (D) = dung. (B) = bone or tooth. 

Garrison No. 1 

Garrison No. 2 

12 230 

± 180 yr B.P. 

13 480 

± 250 yr B.P. 







Sorex sp. 


Ochotona cf. princeps 


B, D 

Sylvihigus idahocnsis 



Thomomys sp. 


Spermophilus sp. 


cf. Peromyscus 


Neotoma sp. 


B, D 

Microtus sp. 



Camelops cf. hesternus 


Ovis or Odocoileiis 


elements. This canyon bottom habitat and possibly the canyon slopes could have sup- 
ported much of the fauna (e.g., Phenacomys and Microtus) recovered from the three 
caves and the rockshelters. Ochotona cf. princeps probably occurred on all suitable 
talus slopes throughout the unglaciated portions of the canyon from the creek level at 
Streamview to Smith Creek Cave. 

Smith Creek Canyon empties into the broad Snake Valley to the east. During part 
of the late Wisconsinan a western arm of Lake Bonneville filled this valley to an 
elevation of 1580 m (Mifflin and Wheat 1979). This high lake stand was only 4 km from 
the entrance of Smith Creek Canyon. The late Pleistocene vegetation of the Snake 
Valley in the area above the lake level is poorly known. 

Thirty km south of Smith Creek Canyon but still within Snake Valley, we re- 
covered packrat middens from a rocky outcrop in the center of the valley (near the 
town of Garrison, Utah, 10 km east of the Snake Range, the closest mountain mass). 
The late Pleistocene age fauna-flora assemblage from the packrat middens is presum- 
ably an indication of the type of habitat above the pluvial lake level but below the 
mountain masses. The isolated rock outcrop provided a suitable habitat for limber pine 
and pika (Table 5; Thompson and Mead 1982). Adjacent areas appear to have been 
a shrub community, with some nearby areas of meadow. Megafauna of the valley 
consisted of at least Camelops cf. hesternus and Ovis or Odocoileus (Table 5). The 
Rancholabrean age fauna of Snake Valley is not well known. A cave acting as a natural 
trap has produced the skull of a wolverine {Gulo gulo), though unfortunately the age 
of the animal is not known (Barker 1976). 

The density of the shrub communities below the mountain mass but above the 
lake level cannot be determined with certainty. The recovery of Sylvilagus idahoensis 
implies that at least some areas were fairly dense with tall stands of Artemisia spp. 
(Hall 1946). Conversely, the recovery of Crotaphytus wisHzeni and Phrynosoma platy- 
rhinos imply areas of relatively open to sparse habitat, possibly exposed playa adjacent 
to the lake. 

Most of the fauna reported here (Tables 3 and 4) come from the inadequately 
dated Reddish-brown Silt Zone in Smith Creek Cave. We have already mentioned that 
the accuracy of the single radiocarbon date on bone fragments from the unit (ca. 28 000 
yr B.P.) is suspect. The minimum age this unit could be is approximately 12 000 yr 
B.P., assuming the stratigraphic associations described by Bryan (1979fl) are correct. 
Equally plausible is that the unit may date from the late Wisconsinan full glacial (ca. 
18 000 to 22 000 yr B.P.) or even middle Wisconsinan (^30 000 yr B.P.). The temporal 
depth of this unit also is not known. Because of these drawbacks we cannot definitely 
state when or what the faunal associations were in the late Pleistocene of Smith Creek 


Canyon. We can state, however, that at least these taxa were in the canyon during the 
late Wisconsinan. This becomes important when considering the species such as Och- 
otona, Phenacomys, and other locally extirpated animals. 

Tanner (1978) has stated that most of the present Great Basin Desert reptiles have 
extended their ranges into the Great Basin as post-Pleistocene introductions from a 
Pleistocene refugium of the southern deserts (Chihuahua, Coahuila, Sonora; Ballinger 
and Tinkle 1972). The late Wisconsinan records oi Crotaphytus wislizeni, Phrynosoma 
platyrhinos, Sceloporus magister, Hypsiglena torquata and Rhinocheilus lecontei in 
the Smith Creek Canyon fauna presented here does not support Tanner's hypothesis. 
If the fauna from the Reddish-brown Silt Zone in Smith Creek Cave is of approximately 
12 000 to 22 000 yr B.P. or even >30 000 yr B.P., then many of the lizards and snakes 
presently inhabiting the eastern Great Basin are not post-Pleistocene invaders. The 
important fact is that desert elements in the modern fauna were already in the Great 
Basin prior to the end of the late Wisconsinan. These results call for reconsideration 
of current biogeographical hypotheses on the evolution of the Great Basin Desert 

At some time in the late Pleistocene and early Holocene, some of the amphibians, 
reptiles, and mammals found in the Snake Range and Snake Valley either adjusted 
their distributions or became extinct. Martin (1967) has previously expressed his model 
that Early Man exterminated the megafauna of North America. The amphibians, rep- 
tiles, and small mammals adapted individually to the climatic changes, however minor 
or major, of the late Pleistocene and Holocene. 

Brown ( 1971 , 1978) and Grayson (1 98 1 , in press a,b) have theorized that local extir- 
pations in the Great Basin of an assemblage of small mammals on a mountain range 
are related to the size of the animal population and the mass of the mountain. In the 
case of the Snake Range (emphasizing Smith Creek Canyon) certain animal species 
became extirpated {Sceloporus magister, Phenacomys cf. intermedins, Ochotona cf. 
princeps, and others) while additional species developed relictual populations, inhab- 
itants of a boreal island in a sea of sagebrush (e.g., Lampropeltis pyromelana and 
Marmota flaviventris). The smaller mammals, at least Ochotona and Phenacomys, 
lingered on into the early Holocene (Grayson 1981; this report). The Ochotona cf. 
princeps remains reported here indicate that suitable habitat and climate occurred in 
the Snake Valley at least until ca. 12 000 yr B.P. and in Smith Creek Canyon until ca. 
6500 yr B.P. (Thompson and Mead 1982). Similar extralocal occurrences of Och- 
otona are documented as late as 7000 yr B.P. in eastern Oregon (Grayson 1981, in 
press). Grayson has argued that the timing of extinction of these relictual small mammal 
(and possibly some amphibian and reptile) populations was in large part determined 
by the size and distribution of habitat islands and by the size of the species population 
on the given habitat island. It is also possible that the mid-Holocene period of warmer- 
than-present temperatures, seen in the elevational raising of upper treeline on the 
Snake Range and elsewhere in the Great Basin (LaMarche 1973, LaMarche and Moo- 
ney 1972) may have reduced the size of montane habitat islands and accelerated the 
rate of extinction of animal populations. 

The faunal account presented here is the first detailed account of amphibians and 
reptiles from late Pleistocene and early Holocene age deposits in the interior Great 
Basin; in addition, the assemblage has provided additional data on mammalian species. 
The Phenacomys cf. intermedins reported here is the first late Pleistocene record of 
the genus in the Great Basin. Because some of the fauna (especially Ochotona) was 
recovered from packrat middens, it is associated directly with a reconstruction of the 
local flora based on plant macrofossils. To our knowledge, similar plant and animal 
associations have not been pursued in this detail for the Great Basin. 

The research in Smith Creek Canyon over the last 50 years has illustrated the 
wealth of information, mainly faunal, available in dry cave deposits of the Great Basin. 
With the recent surge of packrat midden analyses in Smith Creek Canyon, a whole 
new aspect of late Pleistocene and Holocene community reconstruction has emerged. 
Faunal assemblages can now be found in and associated temporally with the wealth of 
flora data recoverable from packrat middens. 



Our sincere thanks goes to Donald K. Grayson for his helpful consultations and 
encouragement in our study of the Great Basin natural history. Gerald R. Smith kindly 
provided the identification of the fish remains and his personal comments. Jeffrey S. 
Green provided the pygmy rabbit fecal pellets for our dung analysis. We also thank 
Susanne Miller and Alan Bryan for allowing us to study the Smith Creek Cave her- 
petofauna from previous excavations, and Donald R. Tuohy for his field assistance and 
guidance. Radiocarbon dates were provided by the University of Arizona, the Uni- 
versity of Waikato, and Geochron Laboratories. Research was funded by National 
Science Foundation Grant DEB75- 13944 to Paul S. Martin. Special thanks are due to 
Marna A. Thompson and Eugene Hattori for helping with field collections, to Emilee 
Mead for the photography, and Deborah Gaines for typing. 

Literature Cited 

Anderson, Elaine. 1974. A survey of the late Pleis- 
tocene and Holocene mammal fauna of Wyo- 
ming. Pages 78-87 in Michael Wilson (editor). 
Applied geology and archaeology: the Holo- 
cene history of Wyoming. Report of Investi- 
gation No. 10. Geological Survey Wyoming, 
Laramie. Wyoming. 

Auffenberg, Walter. 1963. The fossil snakes of 
Florida. Tulane Studies in Zoology 10:131- 

Ballinger, Royce E., and Donald W. Tinkle. 1968. 
A new species of Uta (Sauria: Iguanidae) from 
Salsipuedes Island. Gulf of California, Mexi- 
co. Occasional Papers of the Museum of Zo- 
ology, University of Michigan No. 656. 

Barbour, Roger W., and Wayne H. Davis. 1969. 
Bats of America. University Press of Ken- 
tucky, Lexington. Kentucky. USA. 

Barker, Marcus S. 1976. The wolverine (Gulo lus- 
cus) in Nevada. Southwestern Naturalist 

Brattstrom, Bayard H. 1958. Additions to the 
Pleistocene herpetofauna of Nevada. Herpe- 
tologica 14:36. 

. 1976. A Pleistocene herpetofauna from 

Smith Creek Cave, Nevada. Southern Califor- 
nia Academy of Science Bulletin 75:283-284. 

Brown. James H. 1971. Mammals on mountain- 
tops: nonequilibrium insular biogeography. 
American Naturalist 105:467-^77. 

. 1978. The theory of insular biogeography 

and the distribution of boreal birds and mam- 
mals. Pages 209-227 //; K. T. Harper and 
James L. Reveal (editors). Intermountain Bio- 
geography: a symposium. Great Basin Natu- 
ralist Memoirs No. 2. 

Bryan, Alan L. 1979rt. Smith Creek Cave. Pages 
262-251 (/; Donald R. Tuohy and Doris L. 
Rendall (editors). The archaeology of Smith 
Creek Canyon, eastern Nevada. Nevada State 
Museum. Anthropological Papers No. 17. 

. 1979/7. Council Hall Cave. Pages 2.54-268 

(/( Donald R. Tuohy and Doris L. Rendall (ed- 
itors). The archaeology of Smith Creek Can- 
yon, eastern Nevada. Nevada State Museum. 
Anthropological Papers No. 17. 

Davis. W. B. 1939. The Recent mammals of Idaho. 
Caxton Printers. Limited. Caldwell. Idaho, 

Drewes. Harald. 1958. Structural geology of the 

southern Snake Range, Nevada. Geological 
Society of America Bulletin 69:221-240. 

Goodrich, Robert B. 1965. The Quaternary mam- 
malian microfaunal assemblage of Smith 
Creek Cave, Nevada. Master's thesis. Cali- 
fornia State University at Los Angeles, Cali- 

Grayson, Donald K. 1977. On the holocene history 
of some northern Great Basin lagomorphs. 
Journal of Mammalogy 58:507-513. 

. 1981. A mid-Holocene record for the 

heather vole, Phenacomys cf. intennedius. in 
the central Great Basin and its biogeographic 
significance. Journal of Mammalogy 61:115- 

. in press a. Notes on the history of Great 

Basin mammals during the past 15.000 years. 
//; D. B. Madsen and J. F. OConnell (editors). 
Desert Varnish: a review of anthropological 
archaeology in the Great Basin. Society for 
American Archaeology Memoir 2. 
-. //; press h. The paleontology of Gatecliff 

Shelter small mammals. In D. H. Thomas (ed- 
itor). The archaeology of Gatecliff Shelter and 
Monitor Valley. American Museum of Natural 
History Anthropological Papers, New York, 
New York. 

Gruhn. Ruth. 1979. Excavation in Amy's Shelter, 
eastern Nevada. Pages 90-160 //; Donald R. 
Tuohy and Doris L. Rendall (editors). The ar- 
chaeology of Smith Creek Canyon, eastern 
Nevada. Nevada State Museum. Anthropo- 
logical Papers No. 17. 

Guilday. John E. 1969. Small mammal remains 
from the Wasden site (Owl Cave), Bonneville 
County, Idaho. Tebiwa 12:47-57. 

, and Eleanor K. Adams. 1967. Small mam- 
mal remains from Jaguar Cave, Lemhi Coun- 
ty, Idaho. Tebiwa 10:26-36. 
-. and Paul W. Parmalee. 1972. Quaternary 

periglacial records of voles of the genus Phe- 
ncHomys Merriam (Cricetidae: Rodentia). 
Quaternary Research 2:170-175. 

Guthrie, R. D. 1973. Mummified pika (Ochotona) 
carcass and dung pellets from Pleistocene de- 
posits in interior Alaska. Journal of Mammal- 
ogy 54:970-971. 

Hall, E. Raymond. 1946. Mammals of Nevada. 
University of California Press, Berkeley, Cal- 
ifornia. USA. 


. and Keith R. Kelson. 1959. The mammals 

of North America. The Ronald Press Compa- 
ny. New York, New York. USA. 

Harrington, Mark R. 1934. American horses and 
ancient men in Nevada. The Masterkey, 
Southwest Museum Publication 8:165-169. 

Harris, Arthur H. 1977. Wisconsin age environ- 
ments in the northern Chihuahuan Desert: evi- 
dence from the higher vertebrates. Pages 23- 
51 /// Roland H. Wauer and David H. Riskind 
(editors). Transactions of the symposium on 
the biological resources of the Chihuahuan 
Desert region United States and Mexico. Na- 
tional Park Service Transactions and Proceed- 
ings Series No. 3. 

Hill. William H. 1971. Pleistocene snakes from a 
cave in Kendall County. Texas. The Texas 
Journal of Science 22:209-216. 

Howard. Hildegarde. 1935. A new species of eagle 
from a Quaternary cave deposit in eastern 
Nevada. Condor 37:206-209. 

. 1952. The prehistoric avifauna of Smith 

Creek Cave, Nevada. Bulletin of the Southern 
California Academy of Sciences 51:50-54. 

Jones, J. Knox. Dilford C. Carter, and Hugh H. 
Genoways. 1979. Revised checklist of North 
American mammals north of Mexico. 1979. 
Occasional Papers the Museum, Texas Tech 
University No. 62. 

Kurten. Bjorn, and Elaine Anderson. 1980. Pleis- 
tocene mammals of North America. Columbia 
University Press. New York, New York. USA. 

LaMarche. Valamore C. 1973. Holocene climatic 
variations inferred from treeline fluctuations 
in the White Mountains. California. Quater- 
nary Research 3:632-660. 

. and H. A. Mooney. 1972. Recent climatic 

change and development of the bristlecone 
pine (P. longaeva Bailey) krummholz zone. 
Mt. Washington, Nevada. Arctic and Alpine 
Research 4:61-72. 

Martin, Robert F. 1973. Osteology of North Amer- 
ican Bufo: The Americanus, Cognatus, and 
Boreas species groups. Herpetologica 29:375- 

Martin, Paul S. 1967. Prehistoric overkill. Pages 
75-120 //; Paul S. Martin and Herbert E. 
Wright (editors). Pleistocene extinctions, the 
search for a cause. Yale University Press, 
New Haven, Connecticut, USA. 

Mawby, John E. 1967. Fossil vertebrates of the 
Tule Springs site, Nevada. Pages 105-128 //; 
H. M. Wormington and Dorothy Ellis (edi- 
tors). Pleistocene studies in southern Nevada. 

Nevada State Museum, Anthropological Pa- 
pers No. 13. 

Mifflin, M. D., and M. M. Wheat. 1979. Pluvial 
lakes and estimated fluvial climates of Neva- 
da. Nevada Bureau of Mines and Geology Bul- 
letin 94. 

Miller, Susanne J. 1979. The archaeological fauna 
of four sites in Smith Creek Canyon. Pages 
271-329 //; Donald R. Tuohy and Doris L. 
Rendall (editors). The archaeology of Smith 
Creek Canyon, eastern Nevada. Nevada State 
Museum, Anthropological Papers No. 17. 

Robinson, Michael D., and Thomas R. Van De- 
vender. 1973. Miocene lizards from Wyoming 
and Nebraska. Copeia 1973:698-704. 

Smith. Gerald R. 1978. Biogeography of inter- 
mountain fishes. Pages 17^2 //; K. T. Harper 
and James L. Reveal (editors). Intermountain 
Biogeography: a symposium. Great Basin 
Naturalist Memoirs No. 2. 

. W. L. Stokes, and K. F. Horn. 1968. 

Some late Pleistocene fishes of Lake Bonne- 
ville. Copeia 1968:807-816. 

Stebbins, Robert C. 1966. A field guide to western 
reptiles and amphibians. The Peterson field 
guide series. Houghton Mifflin Company. Bos- 
ton. Massachusetts. USA. 

Stock, Chester. 1936. A new mountain goat from 
the Quaternary of Smith Creek Cave, Nevada. 
Bulletin of the Southern California Academy 
of Sciences 35:149-153. 

Tanner. Wilmer W. 1978. Zoogeography of rep- 
tiles and amphibians in the Intermountain Re- 
gion. Pages 42-53 //; K. T. Harper and James 
L. Reveal (editors). Intermountain Biogeog- 
raphy: a symposium. Great Basin Naturalist 
Memoirs No. 2. 

Thompson. Robert S. 1979. Late Pleistocene and 
Holocene packrat middens from Smith Creek 
Canyon. White Pine County. Nevada. Pages 
361-380 //; Donald R. Tuohy and Doris L. 
Rendall (editors). The archaeology of Smith 
Creek Canyon, eastern Nevada. Nevada State 
Museum. Anthropological Papers No. 17. 

, and Jim I. Mead. 1982. Late Quater- 
nary environments and biogeography in the 
Great Basin. Quaternary Research 17:39-55. 

Valastro. S.. E. Mott Davis, and AlejandraG. Var- 
ela. 1977. University of Texas at Austin ra- 
diocarbon dates XI. Radiocarbon 19:280-325. 

Van Devender, Thomas R., and Jim I. Mead. 1978. 
Eariy Holocene and late Pleistocene amphib- 
ians and reptiles in Sonoran Desert packrat 
middens. Copeia 1978:464-475. 


Volume 20 Number 2 pp. 27-55 25 January 1983 

Fossil decapod crustaceans from the Lower Cretaceous, - ^ " 

Glen Rose Limestone of Central Texas 

Gale A. Bishop 

Department of Geology and Geography, Georgia Southern College. 
Stalesboro, Georgia 30460 USA 

Abstract. The Lower Albian carbonate rocks of the Glen Rose Limestone of Central Texas contain 
a diverse decapod fauna dominated by Protocallianassa species with 10 other taxa represented by fewer 
specimens. The fauna consists of Protocallianassa species, P. klofi, new species, Pagurus handerensis 
Rathbun 1935, Roemerus robustus. new genus and species, Palaeodromites naglei. new species, Diaulax 
roddai. new species, Hillius yoimgi, new genus and species, Dioratiopus scotti, new species, Pseudo- 
necrocarcinus stenzeli, new species, Prehepatus hodgesi, new species, and Torynommal densus, new 


The remains of fossil decapods have been among the most rarely reported fossils 
in the Glen Rose Limestone of Central Texas. Rathbun (1935:35) described Pagurus 
handerensis from a single fragmental specimen. Stenzel (1945:435) revised that descrip- 
tion on the basis of several additional specimens and mentioned that other decapods 
were associated with the P. handerensis remains described by him. "Among the other 
chelae found by Mr. Watkins at the same locality there are some which obviously can 
have nothing to do with Pagurus, because they belong to other well-established and 
unrelated genera such as Callianassa"" (Stenzel 1945:437). 

The fauna described here is important because it testifies to the presence, abun- 
dance, and diversity of the decapod fauna of the Glen Rose Limestone. This fauna 
adds to our knowledge of the Early Cretaceous decapod fauna of North America. A 
diverse fauna of Late Albian age has been described by Rathbun (1935). 

The Glen Rose Limestone 

The Glen Rose Limestone is composed of beds of limestone that are resistant to 
weathering alternating with less resistant marls, giving rise to a characteristic stair-step 
topography (Fig. 1). The presence of mud cracks, bored bedding planes, bedding planes 
with encrusting oysters, algal mat laminations, ripple marks, dinosaur trailways, and 
plant debris is evidence for supratidal, intertidal. or shallow subtidal conditions. Beds 
of lime muds containing echinoids, miliolid foraminifera, infaunal pelecypods, corals 
and numerous gastropods imply marine conditions prevailed at times. The interbedding 
of these sediments points to a depositional system of shallow marine lagoons with 
numerous small islands or rapidly prograding supratidal areas in protected lagoons 
behind a reef-like barrier (Winter 1962, Hendricks and Wilson 1967:5, Stricklin et al. 
1971, Young 1972:1). 

The Glen Rose Limestone of Central Texas contains a scant ammonoid fauna 
which allows correlation with the European section (Young 1972:1 1, 1974:179). The 
Glen Rose Limestone is divided into upper and lower members by a bed (or zone of 
beds) containing numerous steinkems of the bivalve Corhula (Stricklin et al. 1971:23). 


Figure 1. Lithostratigraphy and biostratigraphy of the Glen Rose Limestone (after Young 1974). 

The '"Corbula''' bed is used in this study as a stratigraphic marker to locate the strati- 
graphic position of each collection. All of the Glen Rose Limestone below the Corbula 
bed and to a level of about 41 m (135 ft) above it are considered to be Early Albian 
by Young (1974:176). This includes all of the specimens described in this study. 


Most specimens of decapods have been collected from 3 localities (Fig. 2). The 
specimens are from several collections: (GAB = Gale A. Bishop; SDSNH = San Diego 
Natural History Museum; WSA = W. S. Adkins; UT = University of Texas [Austin]). 

The Nagle Locality (GAB 27).— This exposure was extensively collected by J. S. 
Nagle during the early 1960s. It is situated at the junction of Highways 290 and 281, 
about 9.7 km (6 mi) south of Johnson City, Blanco Co., Texas (Fig. 3). 

Decapods occur throughout about 9 m (30 ft) of Glen Rose Limestone but are 
more abundant in an interval just below a Corbula bed and in a biomicrite about 7.6 
m (25 ft) above the Corbula bed (Fig. 4). 

Interesting assemblages of microscopic claws and dactyli were recovered from 
samples taken at points indicated in the measured section (Fig. 4). The microscopic 
decapod material is particularly abundant in the Salenia texana marl. 

The associated fauna consists of numerous steinkerns of bivalves and gastropods. 

Boerne-Sisterdale Locality {GAB 25). — This exposure is a road cut on Texas Farm 
Road 1376 at the top of a hill (Fig. 5) 12 km (7.5 mi) north of Boerne, Kendall Co., 
Texas. Approximately 9 m (30 ft) of Glen Rose Limestone is exposed in the road cut 
(Fig. 6). The road cut is about 37 m (120 ft) above exposures of the Corbula bed at 
the Hodges Range Section (GAB 26) 1.6 km (1 mi) to the northwest, and probably 


Figure 2. Index map of major collecting localities in Central Texas. 

near the top of the fourth open shelf unit of Young (1974:177). Numerous remains of 
decapods were collected from a biomicrite near the bottom of the road cut (at arrow 
in Fig. 6). 

Decapod remains occur throughout an interval 3 m (10 ft) in thickness but are 
most abundant in a 46 cm ( 1 8 in) bed of fossiliferous biomicrite. The specimens weather 
out of the platey marl and accumulate on the ledge formed by the underlying resistant 

The entire fauna has not been investigated in detail but numerous specimens of 
Homomya are present and preserved in living position. Other elements of the fauna 
include numerous pelecypods, gastropods, and masses of serpulid worm tubes. 


Figure 3. Topographic map of Nagle locality (GAB 21) 9.7 km (6 mi.) south of Johnson City at junction 
of U.S. Highways 290 and 281, Blanco Co., Texas. U.S.G.S. Monument Hill Quadrangle, scale 1:24 000, 
contour interval 20 ft (6.1 m). 





L i me s lo ne 

~^T- Nodular Ls 

Dolomi te 

"I_~_" C I a y s f o n e 

O Decapods 

* Mi crocrustaceans 

I. j_ L I 4^-L__l_-lJ C o r b u I g Bed 

Figure 4. Measured section in Glen Rose Limestone at Nagle locality (GAB 21). 


Figure 5. Topographic map of Boeme-Sisterdale (GAB 25) and Hodges Range (GAB 26) localities, 12.9 
km (8 mi) north of Boeme on Farm Road 1376, Kendall Co., Texas. U.S.G.S. Sisterdale Quadrangle Scale 
1:24 000, contour interval 20 ft (6.1 m). 


Figure 6. Photograph of Boeme-Sisterdale outcrop (GAB 25), decapod interval marked by arrow. Vertical 
scale of foreground approximately 1 cm = 2 m (1" = 17'). 

The Hodges Range Locality {GAB 26).— The Hodges Ranch Locahty (Fig. 5) is 
situated on the third tributary of Wasp Creek west of Texas Farm Road 1376 about 
12.9 km (8 mi) north of Boerne, Kendall Co., Texas. Specimens were collected from 
a soft marl that contained Salenia texana]\xs\ below a Corbula bed (Fig. 1). The locality 
also produced numerous sea urchins and some crinoids. 

The outcrop is a semi-barren gentle slope (Fig. 7) just above a small creek. The 
surface is covered by great numbers of fossils, especially the heart urchin Enallaster 
obliquatus. Other faunal elements include oysters, gastropods, and bivalves. 

The decapod part of the fauna from this locality is almost exclusively dominated 
by claws of the hermit crab, Pagurus banderensis. A collection of gastropod steinkerns 
was made to examine the possibility of preserved pagurids within the lithified mud of 
the steinkerns. No evidence of decapod exoskeleton was observed in the steinkerns. In 
fact, the steinkerns were packed with numerous shells and shell fragments mixed with 
carbonate mud. This suggests a fair amount of washing by currents or bioturbation 
after the shells were last occupied (by hermit crabs?) which would destroy evidence of 
any such occupation. 

Other localities from which Glen Rose decapods have been collected are listed 

1. Bandera-Pipe Creek Road (Texas Highway 16) 1.6 km (1 mi) east of Bandera, 
Bandera Co., Texas; Salenia texana zone. Rathbun 1935:39. 

Figure 7. Photograph of Hodges Range locality (GAB 26). Collecting bag in center 38 cm (1 5 m.) high). 




Figure 8. Line drawings of decapods from the Glen Rose Limestone, Central Texas. A, Palaeodromites 
naglei Bishop; B, Pseudonecrocarcinus stenzeli Bishop; C, Hillius youngi Bishop; D, Dioratiopus scotti Bishop; 
E, Diaulax roddai Bishop; F, Prehepatus hodgesi Bishop; G, Roemems wbustus Bishop; H, Pagurus ban- 
derensis Rathbun 1935; \, Protocallianassa klofi Bishop; J, Torynomma? densus Bishop. 

2. Bandera-Pipe Creek Road (Texas Highway 16) 3.2 km (2 mi) east of Bandera; 
Salenia texana zone. Stenzel 1945:437. 

3. "0.15 mile [0.24 km] south of state Highway No. 29 (Burnet-Austin road) and 
0.07 mile [0.1 1 km] east of the Southern Pacific Railroad track 1.42 miles [2.27 


Table 1. Tabulation of decapod collected from the Glen Rose Limestone. 

GAB-2 1 





Protocallianassa sp." 





P. klofi 




Pagurus banderensi^ 






Roemerus robustus 





Palacodromites naglei 




Diaulax roddai 



Hillius youngi 



Dioratiopus scotti 



Pseudonecrocarcinus stenzeli 



Prehepatus hodgesi 





Torynomma? densus 



" Nearly whole palms. 

'' Numerous other localities yield Protocallianassa. 

" Does not include movable fingers. 


km] airline distance east of the courthouse in Burnet, Burnet Co., Texas." Stenzel 


Junction of Highways 16 and 689, 2.6 km (1.6 mi) east of Bandera; Salenia 

texana zone; collected by Mrs. Henry W. Sebesta, sent to Keith Young by 

George His. 

Blackman Ranch, Little Blanco River, Blanco Co., Texas; WSA 16215 collected 

by A. W. Cleaves. 

Bluff of Blanco River, 4.8 km (3 mi) west of Blanco, Blanco Co., Texas; Salenia 

texana zone 1.5 m (5 ft) below Corbida Bed; UT 45488 (2 specimens) collected 

by N. B. Waechter. 

Shingle Hills Section, Travis Co., Texas; Salenia texana zone; collected by G. 

L. Dawe. 

From a limestone ledge about 1.5 m (5 ft) above water level at Jacob's Well, a 

spring on Cypress Creek, 1.5 km (3.2 mi) northwest of Wimberley, Hays Co., 

Texas, about 55 m ( 1 80 ft) below the Corbula Bed, Lower Glen Rose Limestone 

(calculated from Young 1974). 


The decapod fauna (Fig. 8) of the Glen Rose Limestone consists of 1 1 taxa (Table 
1). Protocallianassa sp. is the most abundant taxon, and Pagurus and Prehepatus are 
second most abundant. The decapod fauna at each locality forms only a part of a much 
larger fauna dominated by molluscs. 

Preservation.— ThQ fossil decapods of the Glen Rose Limestone are found as dis- 
articulated fragments. Claws are most abundant but a few carapaces are present. The 
condition of the remains at the time of final entombment can not be determined. The 
presence of numerous decapod remains is probably due to the similarity of the min- 
eralized exoskeleton and surrounding carbonate rocks. The mineralized exoskeleton is 
often only represented by a chalky carbonate material that weathers readily. This gives 
rise to many steinkerns (internal molds). Steinkerns of brachyurans reflect surface 
morphology but certainly differ significantly from specimens with exoskeletons. The 
finger tips of claws are often filled with sparry calcite signifying that they were not 
completely filled with mud at the time of burial. 

Decapods.— The Glen Rose decapod fauna (Pis. 1-3) gives a crude measure of the 
diversity of decapods in North America in the Early Albian. Because of the small size 
of most of these decapods, they probably are seldom collected and, hence, the antici- 
pated total decapod fauna certainly exceeds that described here. This fauna is the earliest 
Cretaceous decapod fauna described from North America. 


The presence and abundance of hermit crabs (Pagnnis) in these collections is due 
to their size, original abundance in the living fauna, and their heavy mineralization. 
The postulated shallow-lagoonal environment is very compatable with these shallow- 
water decapods. One specimen o^ Pagurus (PI. 2, fig. 17) has an oyster attached to its 
claw. The lack of hermit crab fossils, except claws of pagurids, is due to *the thin 
exoskeleton everywhere but on the heavily mineralized claws that function as an oper- 
culum to close the ''borrowed" gastropod shells in which hermit crabs live. Roemerus 
robustus was also probably a hermit crab. 

Because this is a large, new fauna there are several extensions of geographic range 
of the particularly well-known taxa from the Albian of England (Wright and Collins 
1972). Palaeodromites nag/ei, Diaulax roddai, and Pseudonecrocarcinus stenzeli all 
extend the geographic ranges of the genera from Europe into North America. Tory- 
nomma? densiis extends the range of that genus from Australia to North America. 
Dioratiopus scotti is the second described species of this taxon in North America (the 
other, Dioratiopus dawsonensis (Bishop 1973) is from the Maestrichian of Montana). 
A third species is present on the north flank of the Black Hills, low in the Pierre Shale. 

Hilliiis youngi is so far known from 1 steinkern. It is hoped that additional material 
will substantiate the basic diagnostic features of this poorly represented taxon. 

Prehepatus hodgesi joins other members of the genus, P. cretaceous and P. paw- 
pawensis from the Early Cretaceous of Texas, and P. dilksi from the Late Cretaceous 
Merchantville Fm. of Maple Shade, New Jersey. The pattern and variability of orna- 
menting tubercles on these little claws is extremely interesting. Possibly the variation 
may be due to sexual dimorphism as in the fiddler crabs and perhaps may even have 
had some ritualistic or actual function in this crab's behavior. 

Callianassids. — The most abundant decapod remains found in this collection and 
throughout the Glen Rose Limestone are thalassinids belonging to Protocallianassa, 
Callianassa, Axius, Jaxea, or some other closely related taxon. Only the chelae are 
represented in the collections from the Glen Rose Limestone. 

The thalassinids are differentiated on the basis of carapace morphology, and che- 
lation of walking legs. Within a given taxon (such as Callianassa) sexual dimorphism, 
differing morphology of right and left claws, intraspecific variation, and changes in 
morphology in different instars are all probable (Rathbun 1935:29). These factors seem 
to have combined to yield a baffling spectrum of chelae morphology. The claws found 
in the Glen Rose exhibit a wide variation. Morphotypes can be established easily but 
as the numerous specimens are examined the morphotypes become impossible to 
maintain. The claws will be analyzed again when a graphic analyzing computer system 
becomes available. 

Microcrustaceans. — The presence of crustacean micro fossils in the Glen Rose 
Limestone was pointed out to me by Peter Rodda while curating the collection of the 
Texas Bureau of Economic Geology. Subsequently, microfossil residues were prepared 
from the marls at each locality and numerous other Glen Rose localities. They dem- 
onstrated a diverse and abundant crustacean microfauna consisting of claws and dactyli 
(PI. 3, fig. 44). 

Plate I 

Figures 1-2. Diaulax roddai, 1, Dorsal view of carapace, xl.O; 2, Dorsal view stereo, x2.0 SDSNH 23640 
(orig. GAB 25, specimen 2). Figures 3-5. Pseudonecrocarcinus stenzeli. Dorsal view of latex cast of carapace; 
3, xl.O; 4, stereo, X2.0; 5, view of impression of carapace in limestone, Xl.O, SDSNH 23641 (orig. GAB 
25, specimen 8). Figures 6-7. Dioratiopus scotti, 6, Dorsal view of carapace, Xl.O; 7, Dorsal, stereo, X2.0, 
SDSNH 23642 (orig. GAB 27, specimen 1). Figure 8-1 1. Hillius voungi, 8, Carapace, Xl.O; 9, Carapace, 
stereo, X2.0; 10, Anterior of Carapace X2.0; 1 1, Right side of carapace, X2.0, SDSNH 23643 (orig. GAB 
25, specimen 3). Figures 12-17. Palaeodromites naglei, 2-13, Carapace, Holotype SDSNH 23644 (orig. 
GAB 25, specimen 1); 12, Xl.O; 13, stereo, X2.0; 14-17, Carapace features of specimen SDSNH 23645 (orig. 
GAB 21, specimen 21) collected by Nagle; 14, Dorsal, Xl.O; 15, Dorsal of carapace, x2.0. 


Table 2. Measurements in millimeters of the major claw of Protocallianassa klofi. 


Right or 


Palm length 

Palm height 



















SDNHM 23665 (=25-152) 










































* Specimens in the tables are listed by locality number followed by the specimen number (e.g. 25-67 is 
locality GAB25, specimen 67). 

Systematic Paleontology 

Order Decapoda Latreille 1 803 

Suborder Pleocyemata Burkenroad 1963 

Infraorder Anomura H. Milne-Edwards 1832 

Superfamily Thalassinoidea Latreille 1831 

Family Callianassidae Dana 1852 

Subfamily Protocallianassinae Beurlen 1930 

Genus Protocallianassa Beurlen 1930 

Type species. — Callianassa archiaci A. Milne-Edwards 1860 by original designa- 

Z)/a^«05/5.— "Carapace with linea thalassinica\ first pereiopods with well developed 
chelae, hetereochelous; abdomen with pleura developed on second to sixth somites; 
uropods without diaeresis . . . (Single chelae are hardly distinguishable from those of 
Protaxius or Callianassa).'"' (Glaessner 1969:478). 

Protocallianassa klofi new species 
PI. 3, figs. 41-43; Fig. 81; Tab 2 

Type.—T\yQ holotype, a right major propodus (orig. GAB 25, specimen 152) is 
deposited in the San Diego Natural History Museum (SDNHM 23665). 

Plate 2 

Figures 1-16. Right chelae of Pagurusbanderensis. 1-5, Specimen SDSNH 23646 (orig. GAB 25, specimen 
7). 1, Outer face; 2, Inner face, 3, Bottom, and 4, Top; x2.0. 5, Outer face, Xl.O. 6-8, Specimen SDSNH 

23647 (orig. GAB 2 1 , specimen 1 7). 6. Outer face and 7, Inner face; X2.0; 8, Outer face, X 1 .0. 9-12, Specimen 
UT 45473 (Univ. Texas, Austin). 9, Outer; 1 0, Inner; 1 1 , Lower; and 1 2 Top, X 1 .0. 1 3-1 5, Specimen SDSNH 

23648 (orig, GAB 26, specimen 5). 13, Outer; 14, Inner; and 15, TopX2.0. 16, Specimen WSA 16215 (Univ. 
Texas, Austin), Outer face with walking leg, Xl.O. Figures 17-25. Left chelae of Pagnrus banderensis. 17, 
Specimen SDSNH 23649 (orig. GAB 26, specimen 3), Inner face of left claw with attached oyster, X2.0. 18, 
Specimen SHSNH 23650 (orig. GAB 26, specimen 4). Outer face of propodus, Xl.O. 19, Specimen UT 
45488. Outer face of claw, Xl.O. 20-21, Specimen SDSNH 23651 (orig. GAB 25, specimen 1 1), Outer (20), 
Inner (21), faces of nearly complete minor claw, Xl.O. 22-25, Specimen SDSNH 23652 (orig. GAB 26, 
specimen 1), Outer (22), Inner (23), Bottom (24), and Top (25) views, X2.0. Figures 26-31. Dactyli of 
Pagurus banderensis. 26-29, Specimen SDSNH 23653 (orig. GAB 26, specimen 12), Dactylus of walking 
leg, X2.0. 30-31, Specimen SDSNH 23654 (orig. GAB 26, specimen 1 1), Dactylus of major claw, 30, Side 
Xl.O and 31, Occlusional surface, X2.0. 


Table 3. Height (H), length (L), thickness (T) in millimeters, and height/thickness (H/T) ratios of claws of 
Pagurus banderensis. 








Right Claw 











SDNHM 23647 (= 

















SDNHM 23646 (= 
























SDNHM 23648 (= 












UT 45473 









WSA 16215 

















Left Claw 

SDNHM 23651 (= 

































SDNHM 23652 (= 







SDNHM 23649 (= 










UT 45488 





UT 45488 





* Calculated before rounding thickness measurements from hundreths of a mm. 
** Stenzel's holotype of P. travisensis. 

Plate 3 

Figures 1-19. Prehepatus hodgesi. 1-5, Holotype, Specimen SDSNH 23655 (orig. GAB 25, specimen 14); 
Outer surface of propodus, x 1 .0, 2-5, Outer, Inner, Top, and Distal views of propodus, X2.0. 6-11, Specimen 
SDSNH 23656 (orig. GAB 26, specimen 9); 6, Outer face of propodus, xl.O; 7-1 1, Outer, Inner, Top, Bottom, 
and Distal views, x2.0. 12-13, Specimen SDSNH 23657 (orig. GAB 25, specimen 16); 12, Inner face and 
13, Outer face of propodus, X2.0. 14-16, Specimen SDSNH 23658 (orig. GAB 25, specimen 137), articulated 
carpus and propodus in oblique (14), Front, (15), and Top view (16), X2.0. 17, Specimen SDSNH 23659 
(orig. GAB 2 1 , specimen 22), complete propodus and disarticulated dactylus in front view, X2.0. 1 8, Specimen 
SDSNH 23660 (orig. GAB 25, specimen 18), left propodus in front view, X2.0. 19, Specimen SDSNH 23661 
(orig. GAB 25, specimen 15), left propodus in front view, X2.0. Figures 20-31. Roemems robustus. 20- 
25, Specimen SDSNH 23662 (orig. GAB 26, specimen 8). 20, Outer view, Xl.O. 21-25, Views of outer face, 
inner face, bottom, distal end, and top of left propodus. 26-31, Holotype, Specimen UT 45704. 31, Outer 
view, Xl.O. 26-30, Outer, inner, top and bottom views, X2.0. Figures 32-40. Torynommal densus. 32- 
35, Paratype Specimen SDSNH 23663 (orig, GAB 25, specimen 10), 32, Outer view, Xl.O. 33-35, Outer, 
inner and distal views X2.0. 36^0, Holotype, right propodus. Specimen SDSNH 23664 (orig. GAB 25, 
specimen 5), 36, outer, Xl.O. 37-40, Outer, inner, bottom, and top views, X2.0. Figures 41- 
43. Protocallianassa klofi. Holotype, Right major propodus. Specimen SDSNH 23665 (orig. GAB 25, 
specimen 152). 41-^3, Outer, distal, and inner views, X2.0. Figure 44. Slide of microcrustacean appendage 
elements, X3.25. 
















1 <- 


1 5 


B. Thiclcnes s 





4 0-*- 

1 o — 



^ 30 

20 -« 1 O 

->• 40 

A. Length 

Figure 9. Graph of A. height (mm) vs. length (mm) (bottom) and B. height (mm), vs. thickness (mm) of 
Pagurus banderensis Rathbun 1935. Right claws plotted as dots (read thickness or length on upper scale 
increasing from left to right) and left claws as stars (read thickness of length on lower scale increasing from 
right to left). 

Occurrence, sample size, and preservation. — Fourteen specimens of this taxon were 
collected at GAB 25 and 1 at GAB 21. Most are preserved as single isolated propodi 
with chalky exoskeleton over a firm micrite filling. 

Etymo/og}^. — Named in honor of L. R. KJof, Texas sedimentologist, who often 
exhibited nocturnal and fossorial habits. 

Description. — Fropodus flat, broad, nearly twice as long as high. Palm rectangular, 
slightly longer than high, thin. Upper margin bowed slightly into convex arch. Proximal 
(carpal) edge nearly vertical. Distal margin slants slightly outward to top of short fixed 
finger. Two sinuses present on back of hand along margin; uppermost about Vi the 
distance to top of the fixed finger, the second lies immediately above fixed finger. Lower 
margin convex proximally and concave beneath base of fixed finger. Lower proximal 
corner produced into a rounded projection. Propodus convex on outer face (back of 
hand), nearly flat on inner face (palm). Convexity of outer face continues onto rounded 
upper margin which overhangs inner side (palm) forming a shallow depression along 
top of inner side (palm) parallel to upper margin. Another shallow depression parallels 
the wedge-shaped lower margin on palm. Lower margin of outer side produced into a 
narrow keel from proximal edge almost to base of fixed finger. 

Fixed finger short, nearly horizontal and turned inward. Most large specimens with 
auxiliary ridge along outer edge of occlusional surface of fixed finger, terminating in 
tooth-like projection. An oblique ridge runs ofl' propodus onto fixed finger on back of 
hand (outer side and on palm [inner side]). 


Convex outer face ornamented by 4 or 6 large hair pits along ridge running onto 
finger. Four hair pits arranged in a horizontal row just above level of sinus immediately 
above base of fixed finger. One hair pit situated just above uppermost sinus. Three hair 
pits form broad-based isosceles triangle just below proximal edge of finger ridge. Four 
to 8 hair slits slant upward and distally along lower margin just above fine keel. 

Palm (inner face) has approximately 10 downward and distally slanting hair slits 
along upper margin just beneath overturned angulaled edge where flat palm meets the 
convex outer face. Approximately 15 hair slits slanting upward and distally situated 
along lower edge of palm. 

Comparison.— Protocallianassa k/ofi is similar to P. praecepta Roberts 1962 but 
differs from it by having a relatively shorter palm, a rounded lower proximal corner, 
and lacking the ridge at the base of the fixed finger on the inner face. 

Remarks.— The minor chela of this taxon is not as yet known. No pairs of chelae 
were found preserved together to directly tie the major and minor chelae to one another. 

Protocallianassa sp. 

The majority of the specimens referrable to Protocallianassa comprise a highly 
variable series of chelae. Attempts to differentiate morphotypes failed except in the 
case of P. klofi because gradations were found between all other morphotypes I attempted 
to establish. 

The claws vary from proximally expanded, through rectangular, to nearly oval in 
shape. The cross-sectional shape varies from biconvex, through convex on the other 
face, to spatulate. The fixed fingers are usually curved slightly inward. Ornamentation 
by hair pits is extremely variable. 

Superfamily Paguroidea Latreille 1803 

Family Paguridae Latreille 1802 

Subfamily Pagurinae Latreille 1802 

Genus Pagurus Fabricius 1775 

Type species. — '''Cancer bernhardus Linne 1758" (on official list, ICZN); subse- 
quent designation Latreille 1810 =Eupagurus Brandt 1851 (type. Cancer bernhardus 
Linne' 1758; subsequent designation Stimpson 1858) (obj.).'' (Glaessner 1969:R479). 

Diagnosis. — ""CheWped^ usually dissimilar and unequal, right being much larger 
than left, very rarely subequal; 4th periopods subchelate." (Glaessner 1969:R479). 

Pagurus banderensis Rathbun 1935 
PL 2, figs. 1-31; Figs. 8H, 9; Tab. 3 

Pagurus banderensis Rathbun 1935, p. 39, PI. 9, figs. 7, 8. 

Pagurus banderensis (Rathbun); Stenzel 1945, p. 435, PI. 45, figs. 7-15. 

Palaeopagurus banderensis {Ralhhun); Roberts 1962, p. 175. 

Occurrence. —Specimens of Pagurus banderensis have been collected at many local- 
ities including GAB 21, GAB 25, GAB 26, and Localities 1-7. 

Previous descriptions.— See Rathbun 1935:30 and Stenzel 1945:435-437. 

Remarks. — The collections made at GAB 25 and GAB 26 give the first suites of 
specimens of P. banderensis. Height, length, and thickness data were gathered (Table 
3) and are presented graphically in Fig. 9. 

Size variation is much greater in the right claw than in the left claw. This gener- 
alization also carries over to their morphology and ornamentation; the right claws are 
highly variable in shape, cross section, and degree of granulation, whereas the left claws 
are more consistent in shape and ornamentation. Small right (major) chelae tend to 
have a straighter lower margin which becomes gently convex as size increases. There 
seems to be a tendency for a single row of upper margin granules in small sizes and 2 
distant rows (surrounded by many smaller granules) in larger specimens. 

Two specimens (GAB 21. specimen 17 and GAB 25, specimen 27) are thinner 


Table 4. Measurements in millimeters of claws of Roemerus robustus. 

Total length 


Palm length 







UT 45704 






SDNHM 23662 (=26-8) 
























than the others (H/T-1.62) and might have been called P. travisensis (H/T-2.11) by 
Rathbun. I consider these thinner specimens to be P. banderensis until such time as 
sufficient material becomes available to clearly separate them as P. travisensis upon 
the basis of shape, or until the left (minor) claw of P. travisensis can be demonstrated 
to exist. 

One other specimen (GAB 25, specimen 26) is different enough to merit special 
consideration. The upper margin is narrow and the claw pear-shaped in cross section. 
In front view the lower margin is straight and the upper and lower margins highly 

A walking leg is preserved with a large major chela (PI. 2, fig. 16). A fragment of 
a dactylus of a walking leg is also figured (PI. 2, figs. 26-29). A fairly common element 
in the collections is the movable finger belonging to this taxon (PI. 2, figs. 30-31). 

Family Paguridae Latreille 1802 
Subfamily Uncertain 
Roemerus new genus 

Type species.— Roemerus robustus new species. 

Diagnosis.— ChtXdiQ elongate, similar, with rectangular palm, outer face transversely 
convex, inner flat. Carpal and dactyl articulations perpendicular to lower margin which 
is straight except for convexity below base of fixed finger. Fingers short and turned 
slightly inward. Fixed finger has at least 1 proximal tooth. Tip of movable finger overlaps 
fixed finger in smaller claws and closes onto outer edge of tip in large claws. Surface 
sparsely covered with large, low granules which become numerous and prominent on 
lower edge. The upper surface surmounted by a low, oblique ridge accentuated on the 
proximal inner face by a few low granules and becoming less conspicuous as it runs 
toward the top center of the distal margin of the claw. 

Etymology. — \n honor of Ferdinand Roemer, pioneer geologist and paleontologist 
of Texas and Mexico. 

Comparison.— This claw is similar to Palaeopagurus Van Straelen 1925 but differs 
in being more rectangular, having a more vertical distal margin, and having a slightly 
upturned fixed finger instead of slightly downtumed. Roemerus robustus is easily dis- 
tinguished from Pagurus banderensis Rathbun 1935 by its rectangular shape. 

Roemerus robustus new species 
PI. 3, figs. 20-31; Fig. 8G; Tab. 4 

Types.— ThQ holotype, a left propodus and dactylus (UT 45704), was collected by 
G. L. Dawe from the Salenia texana marl at Shingle Hills, Travis Co., Texas. The 
paratype, a left propodus (orig. GAB 26, specimen 8), was collected at GAB 26. The 
holotype is deposited at the Univ. of Texas (Austin) as UT 45704. The paratype is 
deposited in the San Diego Natural History Museum (SDSNH 23662). 

Occurrence, sample size, and preservation. — ¥'\\q specimens oi Roemerus robustus 
were available for description; 1 from Shingle Hills (UT 45704), 1 from (GAB 26), 
and 3 from (GAB 25). 

Etymology. — The name is taken from the robust nature of the claws of this taxon. 

Description. — Claws robust, similar, rectangular, and thick. Palm slightly longer 




1 n 








5mm 10 15 


Figure 10. Graph of carapace width (mm) vs. length (mm) of Palaeodromites hodgesi Bishop. 

than high and fingers short, slightly incurved, movable finger overlapping tip of fixed 

Lower edge straight except for slight convexity below base of fixed finger. Proximal 
edge perpendicular to lower margin. Distal edge, above the fixed finger vertical. Upper 
margin convex, especially proximally, where it curves down onto prominent carpal 

Outer face transversely convex and ornamented by broad ridges along proximal 
and distal edges formed by narrow furrows on inside. Surface covered by sparse, large, 
subdued granules except for lower edge which has numerous, large, prominent granules. 

Inner face relatively flat and ornamented by a distal ridge and a bend in the 
exoskeleton near proximal margin forming a groove. 

Upper margin surmounted by an oblique ridge which runs from outer-distal corner 
to inner-proximal corner and is progressively more pronounced proximally until it 
forms a noticeable low ridge on upper edge of inner face. A few large granules may 
accentuate ridge. 

At least 1 tooth can be seen situated on fixed finger near its base. 

Comparison.— Roemerus robust us differs from most other pagurids by having a 
pronounced rectangular shape. Only Palaeopagurus Van Straelen 1925 and Petrochirus 
Stimpson 1859 even approach this shape. 

Infraorder Brachyura Latreille 1 803 

Section Dromiacea deHaan 1833 

Superfamily Domiodea deHaan 1833 

Family Dynomenidae Ortmann 1892 

Genus Palaeodromites A. Milne-Edwards 1865 

Type species. — By monoXyxiy', Palaeodromites octodentat us A. Milne-Edwards 1865, 
p. 345, pi. 5; Hauterivian of France. 

Diagnosis.— Carapace broader than long, rounded pentagonal or hexagonal, widest 
two-thirds from front, gently arched transversely and longitudinally. Front square to 
trapezoidal, turned strongly downward; orbits large, oval, widely spaced. Anterolateral 


Table 5. Measurements in millimeters of the carapace of Palaeodromites naglei. 

Length Width 

Specimen (mm) (mm) 

SDNHM 23644 (=25-1) 8.5 10.7 

25^ 9.3+ 11.5+ 

25-52 6.0? 8.6? 

25-53 7.8+ 10.2 

25-54 4.4 5.1 

25-57 9.6+ 11.5 

SDNHM 23645 (=21-21) 12.1 15.6 

borders convex with tooth-like spines or lobes, posterolateral borders short, straight, 
or concave without tooth-like lobes, hind margin short, straight, or concave. Cervical 
furrow clearly defined, sinuous to straight; branchiocardiac furrows weakly defined. 
(After Wright and Collins 1972:49). 

Palaeodromites naglei new species 
PI. 1, figs. 12-17; Figs. 8 A, 10; Tab. 5 

Types.— The holotype SDSNH 23644 (orig. GAB 25, specimen 1) and paratype 
SDSNH 23645 (originally GAB 21, specimen 21) of Palaeodromites naglei are both 
carapaces and are deposited in the San Diego Natural History Museum. 

Occurrence, sample size, and preservation. — Palaeodromites naglei has been found 
at GAB 21 and GAB 25. 

Etymology. — In honor of J. Stuart Nagle, who discovered the first specimen of this 

Description.— Carapace hexagonal, 1.2 times wide as long, very convex longitu- 
dinally, less convex transversely. Carapace furrows poorly developed, a faint cervical 
furrow and 3 branchial furrows present. Rostrum rounded, nearly vertical. Orbits poorly 
defined in dorsal view arching upward. Width between outer angles of orbits 57% 
carapace width. Mesogastric region barely set off from rest of cephalic arch. Three faint 
grooves on branchial region parallel cervical furrow. Metagastric and protogastric regions 
set off from the cardiac and branchial regions by faint muscle attachment scars. Hind 
margin paralleled by a marginal groove. 

Branchial regions split into 4 fields by very faint grooves; anteriormost parallels 
the cervical furrow to a point behind outer angle of orbit then banks backward and 
outward to back of first lateral spine. Second groove beginning at anterior end of muscle 
attachment field, trends outward to first lateral spine. Third furrow extremely faint, 
lying just inside a ridge paralleling posterolateral margin. 

Anterolateral margins each composed of 4 forward facing broad spines which get 
progressively larger posteriorly; the first little more than a broadening of the carapace 
edge just behind outer angle of orbit, the second asymmetrical, bent sightly forward, 
the third the shape of an equilateral triangle, and the fourth broadly rounded, forming 
the widest part of carapace. 

Posterolateral margin serrated by a series of 3 or 4 spines which decrease rapidly 
in size to the last which is little more than a granule, anteriormost a small spine on 
the dorsal shield edge about the size of second anterolateral spine. 

The ventral side and appendages presently unknown because none of these parts 
definitely attached to a carapace. 

Comparison. —Palaeodromites naglei is much smoother and less ornamented than 
other species of this genus. Palaeodromites naglei is additionally distinguished from P. 
sinusosulcatus Wright and Collins 1972 by a straight cervical furrow and lower con- 
vexity, from P. incertiis (Bell 1863) by the lack of coarse granulation, and from P. 
transiens Wright and Collins 1972 by the lack of posterolateral ornamentation. 


Table 6. Carapace measurements in millimeters of Diaulax roddai. 


Length from 

front orbit to 

hind margin 


Orbital wi 






SDNHM 23640 (= 25-2) 




6.1 + 




Family Diaulacidae Wright and Collins 1972 
Genus Diaulax Bell 1863 

Type species. — Diaulax carteriana Bell 1863, by original designation. 

Diagnosis. — ^"Ihe carapace is more or less hexagonal, widest just in front of or 
just behind the ends of the cervical furrow, in longitudinal section curved more or less 
steeply down anteriorly but flat posteriorly, in transverse section flat. The front is 
generally downturned, pointed or squared; it may be sulcate with the edges turned up 
into prominent lobes or nearly flat. The antero- and posterolateral margins are very 
sharp, not lobed and with only a few small, sharp spines directed forwards. The cervical 
and branchiocardiac furrows are weak and tend to be straight and transverse. The 
regions are poorly defined. The surface is very finely granulate." (Wright and Collins 

Diaulax roddai new species 
PI. 1, figs. 1-2; Fig. 8E; Tab. 6 

Type.—T\iQ holotype, a partial carapace (orig. GAB 25, specimen 2) is deposited 
in the San Diego Natural History Museum, SDSNH 23640. 

Occurrence, sample size, and preservation.— Tht holotype is 1 of 3 specimens of 
this taxon thus far collected. It is an almost complete carapace missing only the rostrum 
and left rear corner of the carapace. The exoskeleton is preserved as a chalky limestone 
and does not show surface ornamentation very well. One specimen (GAB 25, specimen 
58) has a partly preserved rostrum and the third specimen (GAB 25, specimen 161) is 
a poorly preserved, crushed carapace. 

Etymology. — ¥ov Peter U. Rodda, Curator of Geology, California Academy of 
Sciences, whose encouragement led to the completion of this study. 

Description. — C2LY?LX)2LCt kite-shaped with truncated anterior and posterior slightly 
longer than wide, widest about '/3 distance from front. Large specimens flat with raised 
anterolateral margins turned slightly under the dorsal shield on pterygostomial regions; 
smaller specimens with higher relief. 

Cervical groove indistinct except for notch where it meets edge of the dorsal shield 
near widest part of carapace. Epimeral muscle scars present but not deeply incised. 

Regions poorly differentiated; rostrum fairly broad, long. Cephalic arch dominated 
by spines at outer edges of orbits and raised anterolateral margins running from orbital 
spines to cervical notch. Gastric areas slightly higher than rest of cephalic arch with 
slight protogastric bosses. Distance between outer edges of orbits -A the carapace width. 
Scapular arch flat, undiflerentiated except for epimeral muscle scars and slight bosses 
distal to epimeral peninsulas, forming ridges that continue almost to hind margin. 

Anterolateral margins dominated by large upward and forward pointing spines at 
outer edge of upward turned orbits that have a single fissure on lower edge, raised 
margin concave to cervical notch. A large upward, forward pointing spine and a second, 
smaller spine lie on margin behind cervical notch (second spine about same distance 
behind notch as orbital spine is ahead of it). Dorsal shield margin convex to first 
scapular spine and runs almost in a straight line to concave V-shaped posterior margin. 
Hind margin bordered by furrow. 


Comparison.— Diaiilax roddai is distinguished from D. oweni (Bell 1850) and D. 
carteriana Bell 1863 by being relatively longer and more flat, especially longitudinally. 
Diaulax roddai most resembles D. feliceps Wright and Collins 1972 but is relatively 
longer, has its maximum width further forward, and has a longer rostrum. 

Remarks. SrmW specimens of this taxon appear to have a better differentiated 
carapace than larger specimens. 

Superfamily Dorippoidea de Haan 1841 

Family Dorippidae de Haan 1841 

Subfamily Dorippinae de Haan 1841 

Hillius new genus 

Type species.— Hillius youngi new species 

Diagnosis.— CdiYapsLCQ pentagonal, fairly flat, slightly wider than long, widest half 
the distance from front. Grooves broad and indistinct. Rostrum broad. Orbital width 
50% carapace width, orbits small, upturned, notched near inner comer, with raised 
rim. Epibranchial areas wide, giving specimens a wing-like or ray-like appearance. 

Etymology. — ¥or Robert Thomas Hill, pioneer Texas geologist, stratigrapher, and 

Comparison.— This taxon has a striking resemblance in carapace shape to Dorippe 
Weber 1795, Goniochele Bell 1858, and Orthopsis Carter 1872. It differs from Dorippe 
by having a wider front with convex anterolateral margins. Hillius differs from Gon- 
iochele by having the widest part of the carapace relatively more forward and formed 
by the epibranchial lobes. Hillius is most similar to Orthopsis but differs from it by its 
having less relief, lack of anterolateral spination, and single orbital lobe. 

Hillius voungi new species 
PI. 1, figs. 8-11; Fig. 8C 

Type.— The Holotype, a carapace steinkern (orig. GAB 25, specimen 3), is depos- 
ited in the San Diego Natural History Museum (SDSNH 23643). 

Occurrence, sample size, and preservation.— The Holotype is the only specimen of 
this taxon's carapace that is nearly complete. The specimen was collected at GAB 25. 
As with any description from decortiated specimens only major ornamentation features 
are likely to be decipherable, and a specimen with exoskeleton will be needed to 
completely define this taxon. 

Etymology. — In honor of Keith Young, Texas Cretaceous stratigrapher and pale- 

Description.— Carapace pentagonal, probably slightly wider than long (partial length 
11.5 mm), widest about half the distance from the front. Carapace slightly convex 
longitudinally; cephalic arch moderately convex transversely, scapular arch fairly flat 
transversely, except at the edges. 

Cervical furrow broad and indistinct except where it crosses dorsal shield margin 
in a pronounced notch continuing on subhepatic region as a well-defined groove to 
base of orbit. An indistinct groove on subhepatic region lies above and is parallel to 
cervical furrow. Faint but distinct mesogastric grooves present. Hepatic grooves broad 
and poorly defined. The most prominent grooves on carapace separate gastrocardiac 
region from branchial regions. Broad grooves separate urogastric, cardiac, and intestinal 
regions, two faint grooves separate branchial regions into 3 parts, the anterior runs 
over carapace edge just behind lateral spine then swings rapidly forward toward cervical 
furrow. Rostrum about 'A carapace width. Orbits small, upturned, with notch near inner 
corner, with raised rim. Distance between outer edges of orbits about 50% carapace 
width. Cephalic arch differentiated into small mesogastric area; large protogastric area 
surmounted by large, low circular bosses; and an upturned hepatic region with at least 
2 small marginal spines immediately ahead of cervical notch. Scapular arch well dif- 
ferentiated into a segmented medial ridge (consisting of urogastric, cardiac, and intes- 
tinal regions) and branchial regions (divided into epibranchial, mesobranchial, and 


metabranchial regions). Urogastric regions have a gentle forward slope and a steep 
posterior slope giving rise to 2 transverse crescentic ridges with small medial spine 
where they meet. Cardiac region diamond-shaped with 2 tubercles symmetrically placed 
across medial axis. Intestinal region poorly defined and partly missing, a single medial 
intestinal tubercle near hind margin. Short longitudinal ridges lie in each gastrobranchial 
groove, perhaps the ridges formed within the epimeral muscle scars. Epibranchial region 
small but forming the prominent lateral wing of this taxon. A small marginal epigastric 
spine lies in posterior part of cervical notch. Two small spines are situated near lateral 
margin of epibranchial wing; the anterior smaller and the posterior larger. Mesobran- 
chial and metabranchial regions with fairly continuous broad longitudinal ridges from 
near the epibranchial groove to hind margin. Posterior of mesobranchial region with 
small boss on this ridge, surmounted by several granules. A small marginal spine 
situated at anterior of metabranchial region. Hind margin missing. 

Comparison.— Hillius youngi differs from Orthopsis bonneyi Carter 1872 by its 
lack of anterolateral spines, single instead of double orbital lobe, more subdued carapace 
relief, and different carapace outline due to the widest point of the carapace being 
farther forward. 

Remarks. — T\\Q line drawing of the carapace of this taxon is based on a single 
steinkem. When further material becomes available the description should be amended 
to include surface ornamentation. 

Family Torynommidae Glaessner 1980 
Genus Dioratiopus Woods 1953 

Dioratiopus Woods 1953, p. 52; Wright and Collins 1972, p. 33, 34, 42. 
Doratiopus Woods, Glaessner 1969, p. 492 (erroneous spelling). 
Glaessneria Wright and Collins 1972 {non Takeda and Miyake 1964), p. 34 ff. 
Glaessnerella Wright and Collins 1975, p. 441. 

Type species.— Homolopsis spinosa (Van Straelen 1936), p. 33; Albian of Valcourt 

Diagnosis. — ''C2iV2ipa.cQ more or less pentagonal with parallel sides, strongly pro- 
jected frontal area, long rostrum with lateral spines and large shallow indistinct orbits 
complete above; the sides are vertical and there are traces of a lateral margin anteriorly, 
but it is normally not sharp or fully developed; the cervical and branchiocardiac furrows 
are strongly marked; a short oblique furrow runs forward from the outer end of the 
branchiocardiac and may extend as far as the cervical, delimiting an epibranchial lobe; 
there is a strong postorbital spine at or just behind the anterolateral angle." (Wright 
and Collins 1972:34). 

Dioratiopus scotti new species 
PI. 1, figs. 6-7; Fig. 8D 

rv'pc — The holotype, a partial carapace steinkern (orig. GAB 27, specimen 1), 
collected at Jacob's Well, Hays Co., Texas, is deposited in the San Diego Natural 
History Museum (SDSNH 23642). 

Occurrence, sample size, and preservation.— The holotype is the only specimen of 
this taxon so far collected. It is decortiated carapace steinkern preserving most of the 
dorsal shield. 

Etymology. — ¥ov Alan J. Scott, Texas Cretaceous and Holocene paleontologist and 

Description.— CsiVdiTp^ce rectangular, longer (partial length 7.9 than wide (partial 
width 6.8 mm). Cephalic arch moderately convex transversely; scapular arch fairly flat 
transversely; carapace relatively level longitudinally. 

Cervical furrow narrow, deep, and prominent; dorsally parallel to anterior dorsal 
shield edge, turning inward and backward, then backward for a short distance cutting 
across medial ridge. Branchiocardiac furrow crosses carapace just behind cervical fur- 


row. Furrows parallel to point where cervical furrow bends inward to cross medial 
ridge, from where branchiocardiac furrow continues backward joining epimeral muscle 
scars and spliting at point near posterior epimeral muscle scar, 1 part swinging inward 
crossing medial ridge as a broad, poorly defined groove and other continuing as epimeral 
muscle scar. Outer arm of epimeral muscle scar loops back inward forming small, flat, 
oval area. Well-defined branchial furrow splits off' epimeral muscle scar near where it 
begins the loop, proceeds outward and forward to dorsal shield edge. 

Rostrum probably triangular, occupying 40% carapace width. Orbits apparently 
small; with sharply upturned rims, occupying 55% carapace width. Mesogastric area 
separated from broad, swollen protogastric regions by shallow, distinct groove, narrow 
anteriorly but rapidly widening at posterior half Protogastric region with small circular 
boss situated at center. A row of 12 small granules begins on raised orbital rim and 
forms an incomplete circle to mesogastric grooves around each protogastric boss. The 
region lying between 2 transverse grooves, crescent-shaped, concave side anterior, with 
short transverse base posterior to middle, giving rise to 2 transverse ridges. Medial 
area behind, where the second groove crosses, a raised region separated by a shallow 
medial groove at its summit into 2 longitudinal ridges. Rear portion of dorsal shield 
missing. Branchial regions divided into 2 parts by branchial cardiac furrows running 
forward and outward; anterior region has single longitudinally expanded granule near 
outer edge, posterior branchial region has 2 granules on outer margin directly behind 
1 on the anterior branchial lobe. The anterior of these 2 is longitudinally expanded 
and large; the posterior round and small. 

Margins of dorsal shield poorly preserved. The photograph, taken before prepa- 
ration was finished, gives the impression of a straight lateral margin (left side) nearly 
to a point on line with the rear of the orbits. The right side appears to be gently convex. 

Comparison.— Dioratiopus scotti is most similar to D. spinosa (Van Straelen 1 936) 
in the size and shape of carapace regions. It differs from D. spinosa by having a wider 
urogastric region with 2 transverse ridges, a longitudinally bilobate cardiac region, 
apparently no furrow delimiting an epigastric region, distal spines on the branchial 
regions, and probably a smoother carapace. The differences in carapace size and shape, 
size, shape, and arrangement of carapace regions, and ornamentation is even greater 
between D. scotti and other congeners. 

Remarks.— The placement of this single specimen into generic level taxon is strongly 
hampered by its mode of preservation as a steinkem and by the obscure nature of the 
lateral margins of the carapace fragment. The discontinuous nature of the lateral margins 
may point to the lack of dorsal pleural sutures, linea homolica, in which case this 
specimen does not belong in Homolopsis. This lack of a straight break and the similarity 
of carapace morphology to Dioratiopus suggests a close alliance with this genus, and 
the specimen is therefore assigned to Dioratiopus until more complete material becomes 

Section Oxystomata H. Milne-Edwards 1834 
Superfamily Calappoidea de Haan 1833 

Family Calappidae de Haan 1833 
Subfamily Necrocarcininae Forster 1968 
Genus Pseudonecrocarcinus Forster 1968 

Type species. — By monotypy; Necrocarcinus quadriscissus Noetling 1881, p. 368, 
pi. 20, fig. 4); Maastrichtian, Limbourg, Holland. 

Diagnosis.— CdiVdiPdiQe wider than long, frontal-orbital margin about Vi carapace 
width. Medial regions poorly differentiated. Mesogastric and protogastric regions com- 
bined into wing-like swellings. Inner side of epibranchial region with ridge, outer side 
with tubercle groups; metabranchial region with weak longitudinal ridges. Sulcus of 
rostrum with 2 or 4 pits. Deeply incised angular grooves form the lateral boundaries 
of the urogastric region. 


Pseudonecrocarcinus stenzeli new species 
PI. 1, figs. 3-5; Fig. 8B 

Types.— The Holotypc, an impression of a carapace, (orig. GAB 25, specimen 8) 
is deposited in the San Diego Natural History Museum (SDSNH 23641). 

Occurrence, sample size, and preservation.— T\\q Holotype, an impression, and 5 
partial carapaces were collected at GAB 25. 

Zinv;;t)/o^v.— Named in honor of Dr. Henryk B. Stenzel, a leader in the study of 
Cretaceous and Tertiary Texas decapods. 

Description.— C^iVSipsiCQ slightly wider than long; widest about V3 distance from 
front. Length from broken tip of rostrum to hind margin 1 1 .8 mm and width (computed 
as 2 times the width of half the crab) 12.5 mm. 

Carapace fairly flat transversely and longitudinally, anterolateral margins lie lower 
than gastric arch. Cervical furrow broad, faint distally but narrower and more distinct 
on central part of carapace near where it ends in a pair of gastric pits; branching at 
distal end, 1 branch continues laterally to dorsal shield edge and other swings anteriorly 
forming a broad, shallow depression which borders anterior side of hepatic region, 
arched orbital region, and arched protogastric region. Epimeral muscle scars deep, 
forming the most noticeable grooves. Inner side of V's thus formed continue forward 
as shallow, broad depressions connecting with cervical furrow; very faint grooves sep- 
arate mesogastric and protogastric regions. 

Carapace regions poorly differentiated by shallow grooves; separation into bosses 
or areoles similarly subdued. Rostrum broad, nearly 25% carapace width. Orbits large, 
arched upward, 2 fissures on rear margins, width between outer edges of orbits about 
50% carapace width. Mesogastric and protogastric regions barely discernable. Two pairs 
of pits lie behind rostrum in mesogastric-protogastric grooves on line with back of 
supraorbital fissures; outermost pair larger, situated slightly more forward than smaller, 
better-defined pair. Protogastric regions with broad, poorly defined bosses. Hepatic 
region with raised boss directly above anterolateral spine. Cardiac regions sharply set 
off' from branchial regions by epimeral muscle scar but barely separated anteriorly 
except for shallow cervical groove, continuing posteriorly with intestinal regions to 
hind margin. Branchial regions broadly arched near epimeral muscle scars, with a raised 
ridge along carapace edge from just behind cervical furrow to a point on line with 
middle part of epimeral muscle scar. 

Anterolateral margin slightly concave for a short distance as it leaves protruding 
outer edge of orbit, dropping in elevation until it begins to become convex, arching 
slightly upward then downward just in front of hepatic spines. Indentations in carapace 
margin in front of hepatic spine and behind it where cervical furrow meets carapace 
edge. Posterolateral margin slightly convex from cervical furrow to hind margin. Hind 
margin concave, bordered by shallow but distinct groove. 

Carapace ornamentation consists of the few broad bosses, anterodistal branchial 
ridges, deep epimeral muscle scar, shallow grooves, and a very fine granulation (slightly 
coarser on medial part) over carapace. 

Comparison. — ThQ genus Pseudonecrocarcinus contains P. quadriscissus (Noetling 
1881). P. biscissus Wright and Collins 1972, and P. stenzeli Bishop new species. All 3 
species have post-rostral pits in common. This character separates this genus from the 
other genera of the Necrocarcininae. In Pseudonecrocarcinus stenzeli and P. quadris- 
cissus the pits are elongated into slits. The carapace of P. stenzeli is much smoother 
than that of P. quadriscissus. 

Remarks.— The presence of pits at the base of the rostrum unite P. quadriscissus, 
P. biscissus, and P. stenzeli into a distinct group. I believe it is best to maintain their 
distinction as a separate genus until such time as we have more data with which to 
judge the phylogenetic affinities of the necrocarcinids. 


Figure 1 1. Tubercle placement on claws of Prehepat us hodgesi Bishop. 

Superfamily Calappoidea de Haan 1833 

Family Calappidae de Haan 1833 

Subfamily Matutinae McLeay 1838 

Genus Prehepatus Rathbun 1935 

Type species. — ^y original designation: Prehepatus cretaceous Rathbun 1935, p. 
47, PI. 11, figs. 29-30. 

Diagnosis. — ChtX^Q small, increasing in height to distal end of palm; fixed finger 
short, movable finger stout; transversely flat to concave on inner face, convex on outer 
face with upper margin broadly rounded to flat and forming an oblique keel which 
overhangs the inner face; surface ornamented by strong tubercles. 

Prehepatus hodgesi new species 
PI. 3, figs. 1-19; Figs. 8F, 11, 12; Tab. 7 

Types.— ThQ holotype o^ Prehepatus hodgesi is a partial right propodus (orig. GAB 
25, specimen 14) deposited in the San Diego Natural History Museum (SDSNH 23655). 
Six paratypes (SDSNH 23656 to 23661) are also deposited in the San Diego Natural 
History Museum. 

Occurrence, sample size, and preservation.— Twenty claws of this taxon are known 
from localities GAB 21, GAB 25, and GAB 26. 

Etymo/ogy. — Named in honor of Floyd Hodges whose fortuitous spilling of coffee 
on his lap caused the discovery of the Boeme-Sisterdale locality (GAB 25) which yielded 
so many fine specimens of this taxon. 

Description.— Carpus with large tubercle on rear distal corner, smaller tubercle on 
front distal corner, 1 on center of dorsal face and 1 midway on rear dorsal margin. 
Margin of proximal, dorsal side with row of 5 granules. 

Right propodus triangular, about twice as long as high (Table 7), palms slightly 
longer than high, highest at distal end of palm. Level upper margin about half the 
length of convex lower margin, flat, horizontally overhanging inner face at proximal 
edge. Carinate proximal part of crest gives way to 4 broad, thin blade-like spines which 
become progressively more vertical distally to base of dactylus. Lower margin trans- 
versely convex. Outer face convex longitudinally, very convex transversely. Inner face 
fairly flat. Two broad, shallow concavities present, 1 below upper margin and 1 at 
proximal end just above lower margin. Propodus ornamented by numerous tubercles 
(situated in 3 lines). Upper sinuous row of 6 tubercles (9, 7, 8, 10, 11) beginning at 
lower carpal articulator follows the carpal margin, swinging forward to follow the upper 
margin to distal edge above dactyl articulator. Second row begins at same point as first 
(near lower carpal articulator), proceeds in sinuous path to base of fixed finger through 
4 large tubercles (counting the first one again) (9, 3, 2, 1) and 1 minor tubercle (18) 
between distal 2 tubercles (1 and 2). The distal 3 tubercles of this row are the most 





T3 p 

-J E 

r- # » # r- 



« + 

I » I » 

» I 


c^- * * c- 

c^- c^- « c^- 

I I 


c^- # I I » ir- * c^- c^- « c^- c^- III 

C-. I # I  I C-. I I I I £-■ C-- C- 

I C-- 

I I 

I I 

I I 

I I 

I I c- 

I I I 

I C-- + « * I * I r- * c^- 

I I I I I * I * 

I « * 

r-»# I !# \ ***** * 

* * * I |#*»»«*»»«C"-» + 

• + 

*  « » 

« « « » 

» « « » 

+ + » » 

00 r-j 
r-' 00 

r- r^ o^ r) o^ (^1 —  

OO \0 i/~l r<"i I^^ ^ <^) 



r~ — Or«-iinoo — r~ 


>n in 00 •^ ■^ vo vO 


O <N NO 
>n r- <N 

rn >/S NO 





1/1 — _ 

lAl r^ 00 On 

od r~-' r-' r-' 











in TtTj-ot^-ONOiNNor-^Tfooo — Tj-voriiri 

(^1 TtONr-Tj-u-iTj-oo — f^iNoo-^inooNOoorj 

ly-^ Tt r^i vd f*-] r^i rr ■^' Tj- -"a- Tt Tt in ly-i H ro nO ■* 

a a 

£ in 

•^ NO 

XNO r^ "T- NO (^J 
r»-) 11 .1. (^ M r^ r-1 m •^ wi so 1^ 00 o^ 

l^'^^'^^lllllllll iin 

C/j (^j (N r^i <N fN r^ fN r) f^i r^i rsi rt n oi n (^i lA 




>— *^ 

m — 

nO <^I 

r^. 00 ■^ 
i^ r) 00 

ri Tt Tt 


p- — m 

On — 00 •^ 

i/-i I/-1 i/-i iy-1 


o o CI :^ NO ^ 

f^J Tj- r^ i ^ (^4 



a o a c. 

nO ON o — 

00 (N C/: 


%i^^i^\i^\i^\irNirNiy-^LJ .minu^uJ iy^»n>/^v" 

^ in >n u^ _ . 
<r- r^ (N r4 1/2 

m in >/S "n 
r^l f^i fN <^J 




<J < 

++ a 












Figure 12. Graph of palm height (mm) vs. palm length (mm) for Prehepatus hodgesi Bishop. Right claws 
plotted as dots (read length on top scale increasing left to right) and left claws as stars (read length on lower 
scale increasing from right to left). 

consistent of the tubercles in presence and relative size. A third row of tubercles (5, 
12, 13, 14) runs in a straight line from a medium tubercle (5) above midpoint of lower 
margin almost to lower edge of the fixed finger. In addition to these "rows" a small 
node is sometimes found in angle formed by the upper rows (16), a medium tubercle 
(6) below middle of upper row, a small one (15) below and behind tubercle 2, a small 
one (17) below tubercle 5, and a small one (19) above and between tubercles 10 and 
1 1 . A spot above tubercle 8 sometimes is roughened or tuberculate. Inner face orna- 
mented by a single tubercle midway along carpal margin. Fixed finger turned downward 
and slightly inward, with faint furrow on lower edge of outer face. At least 2 "teeth" 
are present on prehensile edge; proximal tooth's surface divided into 3 lobes. Movable 
finger strongly curved, with narrow angular ridge on outer side from the articulator to 
midway along dactyl, where it broadens and rounds out to tip. Fingers finely granulate 
and weathered differently than rest of claw, appearing more stable or resistant. 

Left propodus same shape as right, probably also same size (Table 7), outer surface 
with fewer tubercles than right. Tubercle size highly variable, positions more variable 
than those on right claw. A row of 4 tubercles (3, 4, 5, 10) runs in an arc along carpal 
margin from lower articular onto palm. Most other tubercles lie on 2 lines that run 
between major tubercle (1) at base of fixed finger to a large spine (2) distal of lower 
carpal articulator, upper row convex upwards, consisting of 4 tubercles (1, 8, 6, and 
2), lower row convex downward and consisting of 4 tubercles (1,7,9 and 2). Two small 
tubercles (1 1 and 12) may be situated on lower edge of palm below tubercles 1 and 7. 
Three small tubercles (13, 14, and 15) may be present on flattened crest. 

Comparison. — Prehepatus hodgesi has a consistent (though variable) pattern of 
tuberculation different than P. cretaceus Rathbun 1935; P. pawpawensis Rathbun 1935; 
and P. dilksi Roberts 1962. Prehepatus hodgesi is not as rectangular as P. cretaceus 
Rathbun 1935, lacks the great number of spines on the upper margin, does not possess 
the large tubercle near the upper dactyl articulator, and does not have a tuberculate 

Prehepatus hodgesi does not have the numerous fine granules of P. pawpawensis. 


Table 8. Measurements in millimeters of claws of Torvnomma? densus. 


Palm length 



Claw length 

Claw thickness 

SDNHM 23664 (=25-5) 
SDNHM 23663 (=25-10) 





Prehepatiis hodgesi differs from P. dilksi Roberts 1962 in its ornamentation, lack of 
granulate dactylus, and lack of the vertical furrow and rim on the propodus along the 
distal end of the outer surface. 

Remarks. — Tho: taxon shows a surprisingly consistent pattern of ornamentation in 
the arrangement of tubercles and an equally surprising amount of variation in size (or 
presence) of the tubercles. If only a few specimens had been found, it is quite probable 
I would have been tempted to place them in different species level taxa. 

I feel fairly confident that there are at least 2 recognizable instars. The smaller 
specimens are relatively thinner and nearly smooth. A second instar shows tremendous 
variation in tubercle size. A third possible instar may be present and contains the 
largest, most-ornamented specimens. 

The data on cheliped ornamentation are included as I believe there may be a 
behavioral analogy with the claws of fiddler crabs (Crane 1975) and the data may be 
useful to subsequent decapod workers. 

Section Brachyrhyncha Borradaile 1907 
Superfamily Dorippoidea de Haan 1841 

Family Dorippidae de Haan 1841 

Genus Torvnomma V^oods 1953 

Type species. — Torvnomma qiiadrata by original designation. 
Z)/a^A205Z5.— "Carapace subquadrate, widest anteriorly, orbital grooves large, ros- 
trum narrow, oviduct opening on coxa of 3rd periopods." (Glaessner 1969:493.) 

Torvnomma? densus new species 
PI. 3, f^g. 32-40; Fig. 8J; Tab. 8 

Types. — l^YiQ Holotype, a right propodus (GAB 25, specimen 5) and paratype (GAB 
25, specimen 10) are deposited in the San Diego Natural History Museum (SDSNH 
23664 and SDSNH 23663 respectively). 

Occurrence, sample size, and preservation. — Tory nomma? densus is represented 
by 2 right propodi from GAB 25. The holotype is nearly complete except for some 
dissolution of exoskeleton. The paratype is partly crushed on the inner side and has 
the fixed finger broken off Neither specimen preserves the dactylus. The band of dense 
exoskeleton along the distal margin and on the fixed finger is excellently preserved on 
each specimen. 

Etymology. — ¥ov dense exoskeleton along distal edge of claw. 

Description. — Claw nearly twice as long as high, palm slightly longer than high. 
Lower margin slightly convex, tightly rounded at lower proximal corner slanting forward 
along carpal articulation, then broadly rounded to convex upper margin. Distal edge 
nearly vertical concave to base of fixed finger. Fixed finger long, narrow, curving inward; 
decreasing in size distally by 2 steps before it reaches pointed tip. Outer face of claw 
convex, inner face slightly convex. A band of dense exoskeleton present along distal 
edge of claw and on fixed finger. 

Shallow depression on outer face on dense band at base of upper margin of fixed 
finger, giving way to shallow groove that parallels lower margin of finger to its tip. Field 
of small granules near the upper part of dense band and below fixed finger groove. A 
smooth lineation lies on the lower edge of the claw giving rise to an apparent groove. 
Knob-shaped upper carpal articulator granulate. 


Inner face fairly flat, shallow depression below upper margin forming slight over- 
hanging ridge, coarsely granulate on dense band. A shallow, smooth groove running 
along middle of finger joins second step-down. Dense band abundantly granulate below 
groove and sparsely granulate above it. A field of numerous granules lies on the dense 
band even with upper margin of fixed finger. Occlusional surface formed by a broad 
ridge with 2 "teeth" formed where the step-downs in size occur. 

Comparison. — Toryfwmma? densus has a shape similar to Torynomma quadrata 
Woods 1953 but differs by a more convex lower margin, a stouter, stepped-down fixed 
finger, and the band of very dense exoskeleton on the fixed finger and along the distal 
edge of the palm. 

Remarks.— ThQ assignment of this claw to a taxon was most difficult because most 
taxa that have a preserved carapace do not have an adequate description of the chelae. 
The dense exoskeleton on the finger and distal edge of the claw suggests that this may 
be a xanthid-like crab. It was not assigned to the Xanthidae because on xanthids usually 
only the fingers have dense exoskeleton, the fixed finger in xanthids normally has 
"teeth," and this taxon would have extended the range of xanthids from Upper Cre- 
taceous into the Lower Cretaceous. 

It was deemed better in this case to name a new taxon (knowing it will most likely 
be synonomized when the claw is matched to a carapace) than to have yet another 
nameless taxon to refer to in the literature. 


Many persons were directly or indirectly involved in this study. Those who aided 
by helping collect specimens were Nelda Bishop, Susan Deutsch Conger, Arthur Cleaves, 
Lyman Dawe, Tom Grimshaw, Floyd Hodges, Don Lentzen, John Newcomb, Mary 
Beth Bowers Schwartz, and Keith Young. Discussions with Keith Young, H. B. Roberts, 
Alan Scott, and particularly Peter Rodda resulted in a stimulus to complete this study. 
The manuscript was reviewed by Rodney Feldmann and Karl Waage. Jacque Causey 
typed the final manuscript at Georgia Southern College. A Faculty Research Grant 
from the Faculty Research Committee, Georgia Southern College, expedited the com- 
pletion of this work, and an NSF grant (DEB 80115 70) provided time for major revision 
and publication costs. 

Literature cited 

Bell.T. 1850. Notes on the Crustacea of the Chalk 
formation, in F. Dixon (editor) The geology 
and fossils of the Tertiary and Cretaceous for- 
mations of Sussex. Longman, Brown, Green 
and Longmans, London, England. 

Bell, T. 1858. A monograph of the fossil mala- 
costracous Crustacea of Great Britain. Pt. I. 
Crustacea of the London Clay, pages 1^4. 
Palaeontograph. Soc. London. 

Bell, T. 1863. A monograph of the fossil mala- 
costracous Crustacea of Great Britain. Pt. II 
Crustacea of the Gault and Greensand, pages 
1^0. Palaeontograph. Soc. London. 

Carter, J. 1 872. On Oritfiopsis boneyi, a new fossil 
crustacean. Geol. Mag. 9:529-532. 

Crane, J. 1975. Fiddlercrabsofthe world. Prince- 
ton University Press, Princeton, N. J., USA. 

Forster, R. 1968. Paranecrocarcimis lihanoticus 
n. sp. (Decapoda) und die Entwicklung der 
Calappidae in der Kreide. Mitt. Bayer. Staats- 
samml. Palaont. Hist. Geol. 8:167-195. 

Glaessner, M. F. 1 969. Decapoda. Pages 399-65 1 
in R. C. Moore (editor). Treatise on inverte- 
brate paleontology. Geol. Soc. Amer. and Univ. 
Kans. Press, Lawrence, Kansas, USA. 

Hendricks, L., and W. F. Wilson, 1967. Co- 
manchean (Lower Cretaceous) stratigraphy and 
paleontology of Texas. Permian Basin Section 
Soc. Econ. Paleont. Min., Pub. No. 67-8. 

Noetling, F. 1881. Ueber einige Brachyuren aus 
dem Senon von Maestricht und dem Tertiar 
Narddeutschlands. Deutsch. Geol. Gesell. 
Zeitschr. 33:357-371. 

Rathbun, M. J. 1935. Fossil Crustacea of the 
Atlantic and Gulf Coastal Plain. Geol. Soc. Am. 
Special Paper 2. 

Roberts, H. B. 1962. Crustacea. Pages 163-191, 
in H. G. Richards et al. (editors). The Creta- 
ceous fossils of New Jersey: New Jersey Bur. 
Geol. and Top. Paleont. Ser., Bulletin 6 1 , Tren- 
ton, N.J., USA. 

Stenzel, H. B. 1945. Decapod Crustacea from the 
Cretaceous of Texas. Texas Univ., Publ. No. 

Stimpson, W. 1859. Notes on North American 
Crustacea. N.Y. Lye. Nat. Hist., Ann. 7:53. 

Slricklin, F. L., Jr., C. I. Smith, and F. E. Lozo, 
1971. Stratigraphy of Lower Cretaceous Trin- 
ity Deposits of Central Texas. Tex. Bur. Econ. 
Geol. Rept. Invest. 71:1-63. 


Van Straelen, V. 1936a. Contribution a I'etude 
des crustaces decapodes de la periode juras- 
sique. Mem. Acac. r. Belg. CI. Sci. 7:1^62. 

. 1936^. Crustaces decapodes nouveaux 

ou peu connus de Tepoque cretacique. Bull. 
Mus. R. Hist. Nat. Belg. 12(45): 1-50. 

Winter, J. A. 1962. Fredricksburg and Washita 
strata (Subsurface and Lower Cretaceous) 
Southwest Texas. Pages 81-115, in Contrib. 
Geol. So. Tex. The South Tex. Geol. Soc. 

Wright, C. W., and J. S. H. Collins, 1972. British 
Cretaceous Crabs. Palaeontographical Soc. 

Wright, C. W., and J. S. H. Collins. 1975. Glaes- 
snerella (Crustacea Decapoda, Cymonomi- 
dae), A replacement name for Glaessneria 
Wright and Collins, 1972 non Takeda and 
Miyake, 1969. Palaeontology. 18:441. 

Young, K. P. 1972. Cretaceous paleogeography, 
Implications of endemic ammonite faunas. Tex. 
Bur. Econ. Geol. Circ. 72-2:1-13. 

Young, K. P. 1974. Lower Albian and Aptian 
(Cretaceous) Ammonites of Texas. Geoscience 
and Man. 8:1-175-228. 


Volume 20 Number 3 pp. 57-68 24 June 1983 

A new subspecies of Euphyes vestris (Boisduval) from 
southern California (Lepidoptera: Hesperiidae) 

John W. Brown '-'PRAft'Y 

Enloniolog}' Department. San Diego Natural History Museum. 
San Diego^ California 92112 USA 


William VV. McGuire ' 

Saint Francis Hospital. Colorado Springs. Colorado 80903 USA 

Abstract. Euphyes vestris harbisoni. from southern California, is described and illustrated. This 
subspecies is phenotypicalh distinct and geographically isolated from ail other populations oi E. vestris 
(Boisduval). Its range is restricted in part by the distribution of the larval host plant, Carex spissa Bailey 
(Cyperaceae). Life history data are presented including descriptions of the immature stages. Additionally, 
E. ruricola (Boisduval) is removed from the synonymy of £". vestris. 


Euphyes vestris (Boisduval 1852) has long been recognized as a polytypic species, 
with subjective separation of an eastern population, Euphyes vestris Dietacoiuet (Harris 
1862). from the nominate western population. Additionally, it has been appreciated 
by some that a phenotypically distinct population of £". vestris QxisXs in extreme southern 
California (Emmel and Emmel 1973). Data available indicate the presence of this insect 
in scattered populations from Orange County (Orsak 1977) through western San Diego 
County, with speculative distribution to include the northern mountain ranges of Baja 
California Norte. Mexico (MacNeill 1962). However, because sampling and study of 
the southern California population has been extremely limited, definitive analysis had 
not been previously accomplished and its status remained uncertain. Based on the data 
herein presented, it is now obvious that the population of E. vestris centered in San 
Diego County. California, represents a morphologically distinct form that is geograph- 
ically isolated from the nearest E. vestris populations both to the north and east. 
Accordingly, we propose a name for this southwestern population, and report previously 
unknown life history data. 

Depositories abbreviated in the text are as follows: RB. Richard Breedlove. San 
Diego. California: GB. Guy Bruyea. Poway. California: CS. Chuck Sekerman. North 
Hollywood. California: LG. Lee Guidry. Point Loma. California; LACM. Los Angeles 
County Museum of Natural History: SDNHM. San Diego Natural History Museum; 
CIS. California Insect Survey. University of California. Berkeley; and UCI, University 
of California. Irvine. 


Considerable controversy and confusion exist regarding the correct specific name 
of Euphyes vestris (Boisdu\al). The 2 nominal taxa ^ Ilesperia'! vestri.s" [sic] and '"Hes- 
peria ruricola' were described by Boisduval (1852) from specimens collected in Cali- 
fornia by P. Lorquin. The 2 have become confused largely owing to the loss of the type 
of ruricola (fide Oberthiir. Evans 1955); the type of vestris is reportedh in the British 


Museum (Evans 1955). Although ruricola has page priority, the absence of type material 
and the inconclusive nature of the original description have traditionally relegated this 
taxon to junior synonym ol^ vcstris. However, Evans ( 1955) indicates that the description 
of ruricola in "no way" agrees with vcstris. Furthermore, he maintains that the two 
taxa are actually in different genera; ruricola belonging to the genus Hesperia f"abricius, 
and vcstris correctly placed in Euphyes Scudder. This is supported, in part, by the 
question mark placed after the genus ^' Hesperia"' in Boisduval's original description of 
vcstris. The fact that Boisduval questioned the congeneric nature of vcstris and ruricola 
clearly indicates that these two taxa do not represent the same species. Therefore, Evans 
was correct in removing ruricola from the synonymy of vcstris. F. M. Brown (1957) 
states "Holland [(1905)] was responsible for the confusion when he, contrary to most 
of the evidence and the original description, applied [the name] ruricola to the Dun 
Skipper [E. vcstris] ....'' 

Miller and Brown (1981) recently revived E. ruricola "on the basis of a specimen 
that might be the type that is labelled as ^ruricola" in Boisduval's hand . . ." which had 
been located in the collection of the Carnegie Museum. This specimen was designated 
as the lectotype in 1976 by Brown, Miller, and Clench, according to the lectotype label 
(Carnegie Museum lectotype no. 733). This type designation, however, was never 
published, and the senior author's personal examination of the data accompanying the 
specimen revealed a single label that could possibly have been written by Boisduval — 
"Californie (Lorquin)." Contrary to Miller and Brown (1981), there is no evidence to 
indicate that the specimen in question represents Boisduval's concept of ruricola. Since 
a legitimate holotype of vcstris exists in the British Museum (Evans 1955), and the 
description of ruricola is obviously not consistent with vcstris, we propose that ruricola 
be removed from the synonymy of vcstris. 

In agreeing with Stanford (1981) regarding the validity of the subspecific taxon E. 
vestris kiowah (Reakirt 1866). we recognize the following subspecies of £". vcstris: 

Euphyes vestris (Boisduval 1852) 
E. V. v^'^ms (Boisduval 1852) 

i=Pai}iphila osceola Lintner 1878) 

{^Painphila califoruica Mabille 1883) 
E. V. nictaconu't (Harris 1862) 

{^Paiuphila rurca W. H. Edwards 1862) 

{^Hesperia osyka W. H. Edwards 1867) 
E. V. kiowah (Reakirt 1866) 

In addition to the above, the distinctive southern California population of E. vcstris 
deserves formal recognition. Extensive review of the synonymy has failed to uncover 
a previous description that could be unequivocally attributed to this southern California 
population, hence we propose a new name. 

Euphyes vcstris harbisoni new subspecies 
Figures 1-4 

Type material. — \\o\o\ypQ male (SDNHM type L-49), 13.3 km east of Dulzura, 
north slope of Tecate Peak, elevation 500 m, San Diego County, California, 9 June 
1981 (J. Brown): allotype (SDNHM type L-50), 13.3 km east of Dulzura, north slope 
of Tecate Peak, elevation 500 m, San Diego County, California, 14 June 1981 (J. 
Brown). Paratypes: 18 6<5 and 16 99, same locality as holotype, but dates as follows: 1 
3, 1 9, 28 June 1980: 2 53, 2 99, 30 June 1980: 3 <5<5, 1 9, 12 July 1980: 4 99, 9 June 
1981: 4 33, 2 99, 14 June 1981: 2 33, 2 99, 21 June 1981: 1 9, 27 June 1981: 1 9, ex- 
pupa, 2 June 1982: 1 9, ex-pupa, 3 June 1982 (all J. Brown): 6 33, 19, 12 June 1982 
(L. Guidry). 

Disposition c^/O'/^^'-s. — Holotype male and allotype are deposited in the collection 
of the San Diego Natural History Museum: paratypes distributed among collections of 
the following institutions: Los Angeles County Museum of Natural History: California 
Academy of Sciences, San Francisco: United States National Museum of Natural His- 


Figure 1 . Euphyes vestris harbisoni, new subspecies. 6. dorsal. 1 3.3 km east of Dulzura, San Diego County, 

California. 12 June 1980. 

Fr.i RE 2. Kiiphvcs vestris harbisoni. new subspecies, 3, ventral. 13.3 km east of Dulzura. San Diego Countv. 

California, 12 June 1980. 

FiCiURE 3. Euphyes vestris harbisoni, new subspecies, 2, dorsal, 1 3.3 km east of Dulzura. San Diego Countv, 

California, 6 June 1981. 

Fkure 4. Euphyes vestris harbisoni. new subspecies, 2, ventral. 13.3 km east of Dulzura. San Diego County, 
California. 6 June 1981. 

tory. Washington. D. C; Alhn Museum of EntomologN . Sarasota. Florida: Carnegie 
Museum of Natural Histor>. Pittsburgh. Pennsyhania. All remaining paratxpcs to be 
retained by the San Diego Natural History Museum. 

Additionahualerial examined. -CALIFORNIA: SAN DIEGO COUNTY: No fur- 
ther locality. 1 5. 5 June 1936. 1 2, 12 June 1936 (J. Creelman, RB). 1 2. 17 June 1936 
(J. Creelman. LACM). 1 3. 5 June 1936. 1 2. 7 June 1938 (J. Creelman. SDNHM). .San 
Diego. 1 6. 4 July 1920 (O. E. Sette. LACM). San Diego City. 1 <3, 1 2. 14 June 1936 
(J. Creelman, LACM). Adobe Falls. 2 SS. 1 2. 13 June 1936 (F. Thorne. SDNHM). 
Flinn Springs. 1 2. 1 June 1939. 1 c?. 1 5 June 1939. 6 (53. 1 2. 16 June 1939. 2 22. 18 
June 1939 (all W. P. Medlar. SDNHM). 1 3. 18 June 1939 (W. P. Medlar. UCI), 8 <35, 
5 22, 18 June 1939 (W. P. Medlar. LACM). 2 33. 10 June 1980. 1 3. 18 June 1981 (all 
J. Brown, SDNHM). Avocado orchard. Flinn Springs. 1 3. ex-pupa. 15 June 1982 (J. 
Brown. SDNHM). Blossom Valley, 8 km WNW Alpine. 2 66. 3 22. 17 June 1940. 2 
66. 29 June 1942 (all F. Thorne. SDNHM). Hellhole Canyon. 1 3. 12 July 1954 (M. 
Kenney. CIS). Lower Hellhole Creek, 1 3. 1 3 June 1981 (Brown and Brown. SDNHM). 
El Cajon, 2 66, 29 June 1963 (O. Shields. LACM). El Monte Oaks. I 3. 30 May 1965 
(R. Breedlove. RB). 1.6 km W Tccate turn-ofl. 1 3. 30 June 1980(D. Faulkner. SDNHM). 

5 km NW Fallbrook. 1 3. 23 May 1981. 1 3. e.\-pupa. 1 June 1981. 1 2. e.x-larva. 12 
June 1981 (all Brown and Brown. SDNHM). Old Viejas Grade. Poser Mountain. 4 66. 
2 22. 12 June 1981. 2 22. 17 June 1981. 2 33, 26 June 1981 (all J. Brown. SDNHM). 
Behind San Pasqual Academy. 1 3. 30 May 1981 (D. Faulkner. SDNHM). 1 2, ex- 
larva, 30 May 1982 (L. Guidry. LG). Poway Green Valley Truck Trail. 1 3. ex-larva. 

6 June 1982 (L. Guidry. LG). Poway. 1 3, ex-pupa. 15 May 1982 (C. Sekerman. CS). 
Lake Poway. 1 2, ex-pupa, 20 June 1982, 1 6. ex-pupa, 27 June 1982, I 6, 28 June 




Figure 5. 

Figure 6. 

Male genitalia of Euphyes vestris harbisoni. new subspecies. 

Male genitalia of nominate Euphyes vestris. Plantation. Sonoma County, California. 16 May 

1982, 1 9, ex-pupa. 4 July 1982 (all G. Bruyea, GB). ORANGE COUNTY: Silverado 
Canyon, 1 <5. 29 June 1972 (C. Sexton, UCI). 2 <5(5, 8 June 1982 (J. Brown, SDNHM), 
2 <55, 1 9, 5 June 1982 (C. Sekerman, CS). 

Distribution. — Currenlly known only from San Diego and Orange Counties. Cal- 

Diagnosis. — Male: forewing length x= 15.2 mm (range 15.0-16.1 mm). Head, 
thorax and abdomen dark brown, covered with thin light-brown hairs, much lighter 
beneath. Palpi whitish with orange-brown terminal segment. Antennae gold-bi"own, 
ventral surface of club gold, apiculus brown. Forewings chocolate-brown above, with 
lustrous orange over-scaling near a bold black stigma. Stigma composed of two oval 
patches forming an inconspicuously broken black dash. Outer margin of forewing with 
a very fine light tan fringe. Forewings dull brown below, with basal and distal blackening 
representing the undersurface of the stigma. Hindwings same color as forewings above, 




-S 40J 





Z 30 













Figure 7. Flight period graph of Eiiphycs veslris harbisoni. new subspecies, based on wild caught adults, 
using all specimens examined. 

but with less lustrous orange over-scaling; no additional markings. Hindwings dull 
brown below, with no markings. 

Male genitalia: as illustrated (Fig. 5). The structures of the male genitalia of /f. 
vestris harbisoni are very similar to those of both the Arizona populations of i:'. vcstris 
kiowah (not illustrated) and the northern California populations of E. vcstris vcstris 
(Fig. 6). Compared to nominate vcstris. the genitalic capsule of harbisoni is propor- 
tionately longer and slightK less robust, with the vahae more broadh rounded, and 
lacking the ventral caudal indentation commonly present in nominate vcstris. The two 
genitalia figured represent the extremes to more clearh illustrate these differences. 

Female: forewing length x = 16.0 mm (range 14.7-17.0 mm). As in male except 
for primary and secondary sex characters. Forewings dark brown abo\c, with lustrous 
orange over-scaling. Two moderately well-defincd h\aline post-median spots just below 
the discal cell. Females generally with more rounded outer margin of forewing. Fore- 
wings dull brown below, with blackening in basal areas slighth more diffuse than in 
male: the two h\aline spots less well-dcfmed. Hindwings abo\e as in male: dull brown 
below, occasionally with traces of light spots in the discal band area. 

Discussion. — The southern California population of Euphycs vcstris is single- 
brooded, the flight period extending from late May through mid-July (Fig. 7). The 
most distinguishing phenotypic character of the southern California population is its 




Oregon cT 
San Diego Ck). 






<0 001 

Oregon § 
San Diego Co. 





< 0.001 

Arizona (^ 
San Diego Co. 





< 0.001 

Arizona O 
San Diego Co. 





< 0.001 

Arizona (^ 
Oregon (^ 




< O.OIO(NS) 

Arizona ^ 
Oregon § 





< O.IOO(NS) 

Figure 8. Statistical analysis of forewing lengths of various Euphyes vcstris populations using Student's T- 
test comparing two sample means. "P" value represents the probability that the two samples are homogeneous. 
"NS" = not statistically significant. 

larger size relative lo populations from northern California. Oregon and Washington. 
Its larger size and extensive orange over-scaling clearly distinguish it from the Rocky 
Mountain E. vestris populations, which have been referred to as E. vestris kiowah by 
some authors (Stanford 1981), and the eastern E. vestris nietaconiet. There are some 
populations of E. vestris in Arizona that may be referable to E. vestris kiowah. but at 
this time have not been thoroughly studied. Again, these differ from the southern 
California population in both size and maculation. An analysis of the male and female 
forewing measurements between various populations sampled (Fig. 8) demonstrates 
the statistically significant difference between the San Diego County population and 
others based on this parameter. 

The habitat occupied by the southern California population is remarkably dissim- 
ilar to that of all other known populations of £■. vestris. The southern California pop- 
ulation is extremely local, generally occurring in chaparral or riparian communities in 
narrow canyons where there is a seep or spring providing perennial water (Figs. 9- 
12): populations found to date have been at elevations below 650 m. The o\iposition 
substrate and lar\ al hostplant is Carex spissa Bailey (C\ peraceae), with which the adults 
are commonly associated (Fig. 13). As indicated by figure 14. C. spissa has a limited 
and spotty distribution from San Luis Obispo County. California, to Baja California 
Norte. Mexico (Munz 1974). The range of the host plant is obviously a factor limiting 
the distribution of the butterflN. as supported by the fact that all known populations 
of E. vestris liarlusoni (Fig. 15) fall within the range of C. spissa. Howc\er. factors 
other than host plant availability must contribute to restrict the insect's range, as 
indicated by the fact that the butterfly occurs within only a small portion of the host 
plant's distribution. 

Initial observations and efforts to study this insect centered on examination of 
previously captured material and old collection records. Howcxer, there were few 
previous records for the southern California population of Ii. vestris. and the last 
specimen collected prior to this study was in 1 969 (Stanford /// litt.). Many prior records 
came from the Adobe Falls area near San Diego State University, but current devel- 
opment in this area has altered the habitat to the point where it no longer supports the 

63 9. Typical chaparral habitat near Tccatc Peak. San Diego County. C'alilbrnia. 

Fk.iki 10. Habitat near Dul/ura. Caiirornia. 

Fu.i HI 11. Seep area at Dul/ura k)cality. 

FicL HI 12. Habitat near Dul/ura. California. 

Fk.iki 13. Larval host C'(/'c\ s/j/sv^/ HailcN. 

insect. The only fccent coUcclions outside of San Diego C"ouni\ were made \n the 
Silverado Canyon area of Orange County (Orsak 1977). 

Using prior collection records as a guide, we examined iikeh habitats in 1980 
and 1981 in San Diego County and I3aja California Norte. .Adult specimens were found 
first near Flinn Springs Count\ Park, and additional specimens later found in other 
localities noted abo\ e. Fourth instar lar\ ae were subsequentK observed 1 3.3 km east 



San Francisco 

FiC.URE 14. 


Approximate distribution of Carex spissa in southern California and adjacent Baja California, 

of Dulzura in January 198 1. All larvae were found in typical hibernacula formed from 
C. spissa (Fig. 16). These hibernacula were formed by the attachment of two. three or 
four leaves creating a silk-lined, tube-like chamber 7-12 cm long and open at the 
superior end. The chamber itself is usually located in the superior one-third of the 
Carc.x leaves. Each leaf or blade is characteristically bent, generally at less than a 100° 
angle, allowing them to be pulled together and joined with silk by the larva. This type 
of hibernaculum is fairly characteristic and consistent with previous observations made 
on other members of this genus, including /:. a/aha mac {Lindsey 1923). /:. macguirei 
(Freeman 1975). and /:. ciukcsi (Lindsey 1923) (William McGuire personal observa- 

Jjynio/ogy.— The subspecies /:. vestris harhisoni is named in honor of Charles 
Harbison, Curator Emeritus. San Diego Natural History Museum. Mr. Harbison has 
been a patient instructor, and enthusiastic field entomologist, and a pioneer in inves- 
tigating the entomofauna of Baja Califorinia, Mexico. 



mvEnsiDE CO. 



HFIinn Springs 

•• Tecate 

FiCii'RF 15. Distribution of Euphyes vcstris harhisoiii. new subspecies. 

Life History 

Early stages. — The early stages of E. vestris harbisoui closely resemble those de- 
scribed by Heitzman (1964) for the eastern E. vestris metacomet. The larvae of E. 
vestris harbisoni, however, are much larger, and the pupae dark brown rather than 

Egg. — (Fig. 17). 1.5 mm. hemispherical, pale yellow-green with an irreg- 
ular red splotch on the apex. An irregular red band encircling the entire egg midway 
between the base and the apex. 

Eirst inslar. — Length 7.0 mm. Head: shiny brown. Body: ground color translucent 
green, abdominal segments 9 and 10 translucent yellow-tan. Several long, fme hairs 
projecting upwards from the last body segment: thin black collar just behind the head 
indicating the prothoracic shield. 

Second inslar. — LengXh 14.0 mm. Head: yellow-orange with a conspicuous cara- 
mel-colored band encircling the head along its outer margins except above the man- 
dibles. Mandibles dark brown. An oblong dark brown spot set vertically in the upper 
center of the face. Body: ground color translucent green, abdominal segments 9 and 10 
translucent, revealing yellow-tan internal organs. Body co\ered with numerous white 
setae, several longer thin hairs at end of the last segment. Each spiracle indicated by a 
fine black dot. 

Third instar. — Lcnglh 21.0 mm. Head: cream-colored with a well-defined dark 
caramel-colored longitudinal stripe on each side of the face, and a light brown band 
extending from the dark spot to the mandibles. Body: ground color translucent green, 
abdominal segments 9 and 10 more translucent; covered with fine white hairs. Spiracles 
indicated bv a fine black dot. 


Figure 16. Typical hibernaculum formed from Carex spissa. Arrow indicates position of the larva. 

Fourth />75/(3r — Length 28.0 mm. Head: caramel-brown with an oblong black spot 
in the upper center of the face; a narrow cream-colored longitudinal band running 
parallel to, and on each side of the black spot; a cream-colored band around the outer 
edge of the jaws. Body: ground color green, with numerous thin white dashes of in- 
consistent length (previously visible only under microscopy). 

Final instar. — (Fig. 18). Length 36.0 mm. Head: caramel-brown with two cream- 
colored vertical bands; a black oval spot in the upper center of the face. Body: green 
with extremely thin, wavy longitudinal white dashes. A single, subtle, darker green 
longitudinal stripe along the center of the dorsal surface. Minute black dots sprinkled 
over entire body. Terminal abdominal segment translucent pale green. Prothorax white 
with prothoracic shield indicated by a thin black line running into the enlarged first 
spiracle dot, forming a thin black wedge. Each spiracle indicated by a black dot, larger 
for the first and anal spiracles. Ventral surface of abdominal segments 8 and 9 covered 
with a fine layer of silver-white cottony material. 

Pupa. — (Fig. 19). Length 25.0 mm. Dark brown. Abdomen slightly dusted with 
white cottony material. Most of body covered with short, fine bristles; wing cases bare. 
Bristles on head and end of abdomen longer. Tongue case long, extending beyond the 
fourth abdominal segment, lying along the mid-dorsal surface of the pupa, detached 
above the abdominal segments. 

Behavior. — Eggs are laid singly, usually on the undersurface of the leaf blades near 
the base of the plant. During the first instar the young larvae are generally found along 
the mid-ridge crease on the upper surface of the leaves, near the base of the plant. The 
second and third instar larvae, when not feeding, are usually found in silk-lined tubes 
constructed by the larvae by attaching two to four leaves together longitudinally. It is 
during the fourth, or occasionally the third instar, that the larvae construct the over- 
wintering hibernacula previously described, and enter a short diapause. The head is 
always aimed towards the superior end of the hibernacula. 

During the last instar the larva consumes the portion of the leaf blades extending 
beyond the superior end of this hibernaculum. This tube is then abandoned, and the 
larva moves to a dilTcrent location on the plant, or to a different plant and constructs 
a new chamber. The upper portion of these leaf blades are also usually eaten by the 



FiGi'RE 17. Egg of Euphyes vestrls harbisoni. new subspecies. 

Ficii RE 18. Last instar larva of E up fives vestris harbisoni. new subspecies. 

Fita RE 19. Pupa of Euphyes vestris harbisoni, new subspecies. 

larva. When a larva is ready to pupate, it fills the upper end of the tube with a flocky. 
cottony plug 8-10 mm thick. The larva is generally situated in the tube with the head 
towards the superior end: the pupa is always situated in this manner. Pupation lasts 
18 to 21 days. 

Adults emerge from late May through June, the flight period extending into early 
July. At one locality a last instar larva, a pupa, and a single adult were collected on the 
same day (23 May 1981). 

Males commonly ''patrol" the canyons, never venturing far from the vicinit\ of 
the host plant. Both sexes are avidly attracted to nectar sources which include morning 
glory {Calystegia macrostegia tcnuifolia (Abrams) Brummitt), red thistle (Clrsiuni oc- 
cidentale (Null.) Jepson). loosestrife (Lythrum californicwn T. & G.), and rarely, golden 
yarrow {Eriophyllum confcrlifloruni (D. C.) Gray) and black mustard (Brussica nigra 
(L.) Kock.). Commonly females are observed perching on the C'arc.x and basking in 
the late morning and early afternoon sun. 


Euphyes vestris harbisoni is isolated by geograph\ and by the availability of a select 
host plant. While several of the populations found appear to be stable, all are extraor- 
dinarily small. Some of the prior known colonies have been extirpated by construction 


projects and other habitat modifications. A search for new populations and extension 
of range should appropriately be directed into northern Baja California and northward 
up to and including San Luis Obispo County, thus encompassing the known range of 
the larval host plant. Carex spissa. This search in the projected northern range of the 
insect is important when considering potential intergradations with the nofthern Cal- 
ifornia E. vestris vestris. E. vesths harbisoni appears to be morphologically and bio- 
logically distinct, with significant geographic separation from all other known E. vestris 
populations. This restricted range, coupled with the continued threat of habitat deg- 
radation as has been demonstrated in the immediate San Diego city area, makes it 
imperative that careful monitoring be continued. 


We are most grateful to the following for providing assistance and encouragement 
during the course of this study: David Faulkner, Dr. Reid Moran. Dr. Ray Stanford. 
Dr. John Emmel. and Dr. Amadeo Rea. Special thanks go to Poody Brown. William 
Smithey, and John Johnson, all of whom provided photographic assistance, and to 
Marie HoflT Steinauer. and Shirley Latislaw for providing the maps and graphs. 

We would also like to thank the following for providing specimens and/or collecting 
records: Richard Breedlove. San Diego. California; Fred Thorne. Curator Emeritus, 
San Diego Natural History Museum; Julian Donahue, Entomology Section, Los Angeles 
County Museum; Guy Bruyea. Poway. California; Lee Guidry. Point Loma, California; 
Chuck Sekerman. North Hollywood, California; and Gordon Marsh. University of 
California, Irvine. Additionally, we would like to thank Dr. Ginter Ekis. Carnegie 
Museum, for the loan of the unpublished supposed lectotype of Euphycs rwicola. 

Finally, we would like to acknowledge the Xerces Society for their financial support 
of this project, and their recognition of the sensitive status of this insect. Their support 
allowed one of us (John W. Brown) to undertake the extensive field work which made 
this study possible. 

Literature Cited 

Boisduval, J. A. 1852. Lepidopteres de la Cali- 
fornie. Ann. Soc. Ent. France, ser. 2, 10:315, 

Brown. F. M. 1957. Colorado butterflies, part 5. 
Proc. Denver Mus. Nat. Hist. 7:313. 

Emmel. T. C. and J. F. Emmel. 1973. Butterflies 
of southern California. Nat. Hist. Mus. Los 
Angeles Co.. Sci. Ser. 26:1-148. 

Evans. W. H. 1955. A Catalogue of the American 
Hesperiidae in the British Museum (Natural 
History), part 4. London, Trustees British Mu- 
seum. 499 pp. 

Heitzman. R. J. 1964. The early stages of Ew/?/?;^^^ 
vestris. J. Res. Lepid. 3(3): 15 1-1 53. 

MacNeill, C. D. 1962. A preliminary report on 
the Hesperiidae of Baja California. Proc. Calif. 
Acad. Sci. 30(5):91-1 16. 

Miller. L.D. and F. M.Brown. 1981. A catalogue/ 
checklist of the butterflies of America north of 
Mexico. Lepid. Soc. Mem. 2:1-280. 

Munz, P. A. 1974. A Flora of Southern California. 
Univ. Calif Press, Berkeley. Los Angeles, Lon- 

Orsak, L. J. 1977. Butterflies of Orange County. 
Univ. Calif Irvine. Mus. Syst. Biol. Res. 
ser. 4. 

Stanford. R. E. 1981. //; Ferris, C. D.. and F. M. 
Brown (editors). Butterflies of the Rocky 
Mountain States. Univ. Oklahoma Press, Nor- 


Volume 20 Number 4 pp. 69-79 24 June 1983 

Two new idoteid isopods from Baja California and the Gulf of 
California (Mexico) and an analysis of the evolutionary history 
the genus Colidotea (Crustacea: Isopoda: Idoteidae) fARY 

Richard C. Brusca 

Department of Marine Invertebrates, San Diego Natural History Museum. 
San Diego, California 92112 iSA 

Abstract. Two new species of idoteine isopod crustaceans are described from Baja California, 
Colidotea wallersteini new species and Synidoteafrancesac new species (Crustacea: Val\ifera: Idoteidae: 
Idoteinae). The genus Colidotea is rediagnosed. a key to the known species presented, and its evolution 
and historical biogeography described. Both trans-oceanic dispersal and vicariance phenomona appear 
to have played roles in the establishment of modern distributional patterns of Colidotea. 


The idoteid isopod fauna of the northeast Pacific has. in the past 10 years, come 
to be quite well known (see Brusca and Wallerstein 1979b for an introduction to the 
literature). The present study adds 2 new species to this fauna, based on material 
collected by Dr. D. G. Lindquist (University of North Carolina) and some previously 
unsorted samples of microcrustaceans at the Allan Hancock Foundation (University 
of Southern California). 

The terminology used in this paper is standard for the idoteid isopod literature 
(see Menzies 1950 for a summary). The following abbrev iations arc used: AHF. Allan 
Hancock Foundation; USNM, National Museum of Natural History; SDNHM. San 
Diego Natural History Museum. 


Order Isopoda 

Suborder Valvifera 

Family Idoteidae 

Subfamily Idoteinae 

Genus Synidotca Harger, 1878 

The systematics and biogeography of the genus Synidotca were reviewed by Men- 
zies and Miller (1972). who recognized 36 valid species and subspecies. 8 of which 
occur in California. Iverson (1972) described a ninth species from California {S. Dwdia). 
Brusca and Wallerstein (1979a) presented new distributional records for S. harfordl 
Benedict, 1897, which at that time was believed to be the southernmost ranging species 
of the genus in the northeastern Pacific. They noted its disjunct occurrence: central 
California to Magdalena Bay (SW Baja California), Mexico, reappearing again m the 
central Gulf of California. Mexico (2 known records). Brusca and Wallerstein (1979b) 
reviewed the distributions of the 7 known northeastern Pacific shallow-water Synidotca. 
noting that all but S. harfordi were entirely restricted to the cold-temperate waters of 
the Arctic. Aleutian and Oregonian Provinces. Brusca (in press) discusses phylogenetic 


Figure 1. Synidotea francesae new species. Holotype, AHF 736. Female. 

relationships, evolutionary history and zoogeography of Synidotea. as well as the 21 
other genera of Idoteinae. 

Synidotea francesae new species 
Figures 1, 2. 3 

Types. — HoloXype: female. AHF 736; Allotype: male. AHF 736a. Paratypes: 1 
female, SDNHM; 1 female, USNM. 

Locality. — AW specimens from single collection: Mexico, Gulf of California, So- 
nora. El Golfo dc Santa Clara (about 4 miles SE of town); found on sandy beach, 
"scavenging" on cast up tubes of Chaetopterus species (Polychaeta); air temperature 
24°C; water temperature (surO 19°C; 17 April 1973; collected by D. G. Lindquist. 

Diagnosis. — CQp\\3.\or\ without horns or other projections; frontal margin convex; 
eyes bulge outward; body without tubercles or rugae; plcotclson wider than long; per- 


FidiRF 2. Synlciotca francesac ncv,' species. Holotype. AHF 736. Female, a. antenna 1. b. antenna 2. c, left 
mandible, d, lacinia of right mandible, e, maxilla 1. f. maxilla 2. g. maxilliped. h. pereopod I. i. pereiopod 
IV. J. pereopod VII. k. uropod. 


Figure 3. Synidotea francesae new species, a, pleopod I (holotype). b, pleopod 2 (holotype). c, appendix 
masculinum of pleopod 2. male (allotype), d, pleopod 5 (holotype). 

eonites V-VII manifestly shorter than I-IV; median dorsal pattern rounded, as in S. 
laticauda; dorsum with heavy pigmentation. 

Description. — hQxxgXh to at least 6 mm; body ovate and darkly pigmented: dorsum 

Cephalon: Lacking horns or other projections; frontal margin convex, although 
anterolateral angles somewhat expanded; eyes elevated, on distinct bulges. Antenna 1 
with tri-articulate peduncle and uniarticulate flagellum; distal third of terminal article 
with simple setae and esthetascs. Antenna 2 with 5-articulate peduncle and 8-articulate 
flagellum; all articles with setae; articles 4 and 5 of peduncle each with distinct distal 
patch of chromatophores; flagellum quite short, extended only to pereonitc II. Max- 
illiped typical for the genus, with single coupling hook. Maxilla 1 typical for the genus; 
inner lobe with 2 stout plumose setae; outer lobe with 6 long simple setae and 4 short 
pectinate setae. Maxilla 2 typical for the genus, with plumose, simple and comb setae 
as figured. Mandible with 4-toothed incisor and large molar process, the latter smooth 
on outer margin and toothed on inner margin; lacinia mobilis of left mandible 3- 
toothcd; lacinia of right mandible 3-toothed but with additional large serrate spinelike 

Pereon; Manifestly elevated along middorsal line; entire dorsal surface with dark 
pigmentation, except along lateral margins. Pereonites I-IV large, long and without 
dorsal coxal plates; pereonites V-VII small, short, and with large dorsal coxal plates, 
visible in dorsal aspect by presence of faint suture lines just median of edge of chro- 
matophorc pattern; all pereopods form ventral coxal plates that meet in midline of 
sterna. Median dorsal pattern of pereonites II-IV {scusu Menzies and Miller 1972) with 
rounded posterior margins. Pereonites I V-VII slightly produced along posteromedial 
margin. Pereopod I with dactyl nearly as long as propodus. 

Picon; Posterior border with faint indentation, otherwise evenly convex. Pleon 
(=plcotelson) width equal to length, or up to 1.5 times wider than long. Plcopods 1-3 


Table 1. Principal attributes distinguishing Colidotca and its sister-group. Synisoma. from each other and 
from all other genera of Idoteinae.* 



Flagella of second an- 




Number of articles in 
maxillipedal palp 

Appendix masculina (of 


long, arising from base of endo- 
pod (of second pleopod) 


long, arising from base of endopod 
(of second pleopod) 

Lacinia mobilis 

very large; as large or nearly as 
large as incisor process 

very large; as large or nearly as large 
as incisor process 


relatively short: much less than 
'A total body length 

relatively long; 'A or more total 
body length 

Pleomere fusion 

all pleomeres fused; one pair lat- 

all pleomeres fused; no lateral inci- 

eral incisions present 

sions remain 


very small (except in C. findleyi); 
set on lateral margins of cepha- 

very small; set on lateral margins of 

For comparable data on the other 20 genera of Idoteinae see Brusca (in press). 

with plumose marginal setae, decreasing posteriorward; pleopods 4-5 without plumose 
marginal setae. Uropod with 3 stout plumose setae at junction of protopod and lamella. 
Appendix masculinum of male simple, with weakly grooved inner margin and a few 
distal setae; penes entirely fused into large tongue-like flap. 

Ety}}iology\ — Synidotea fraucesae is named in honor of Frances Runyan: biological 
illustrator, horticulturist, and friend. 

Discussion.— Synidoicajrancesae is similar to S. laticaucia Benedict, 1897, and S. 
harfordi Benedict. 1897 (both known from California shoi"es), in having a smooth 
dorsum with a characteristic medial dorsal pattern, evenly convex lateral margins on 
the body somites, elevated (bulging) eyes, and notch at the apex of an otherwise evenly 
convex pleotelson. It differs most strikingly from these species in having a pleon as 
wide or wider than long, a barely perceptable notch at the pleotelson apex (rather than 
a deep notch), pereonites V-VII manifestly reduced in size, and a distinct dorsal chro- 
matophore pattern. It differs further from 5". laticauda in having an evenly convex 
tVontal margin, a 4-toothed incisor on the mandible, and shorter antennae. It differs 
further from S. harfordi in having rounded posterior margins on the medial dorsal 
pattern of pereonites II-IV, shorter antennae, and a coupling hook on the maxillipedal 

Of the 18 species o^ Synidotea now known from the eastern Pacific, only S. fran- 
ccsac is a regular resident of warm waters (i.e., the Eastern Pacific Zoogeographic 
Region); all others are temperate or polar species. The genus is worldwide in distribution 
but notably absent from the New World tropics. Brusca and Wallerstein (1979b) and 
Wallerstein and Brusca (1982) hypothesized that this absence is due to the inability of 
Synidotea species to develop certain morphological (size; body spination) and life 
history (early reproduction; predator avoidance behaviors) adaptations that have al- 
lowed successful radiation in this region by other genera (e.g., Erichsoiwlla. Eusyni- 
nicriis, FarasyniDicrus). and that any species of Idoteidae that may be found successfully 
inhabiting the warm coastal waters of the tropical eastern Pacific would have had to 
evolve some of these adaptations. In the subtropical waters of the Gulf of California. 
S.francesae has evolved at least 1 (small size) if not more (e.g., early reproduction) of 
these attributes which presumably ha\e played a role in its ability to survive the 
predation-intense summers of this region. Menzies and Miller (1972) state that the 
mean length of the 8 Arctic species ol\Synidolca is 18.8 ± 2.6 mm, while the mean 
length of the 5 tropical species then known (all from the Old World) was 8.0 ± 2.2 
mm. Temperate species fall between these two extremes. By comparison, the largest 
specimen oWSynidoteafrancesae examined is onh 6 mm in length. 


Genus Colidotea Richardson, 1905 

The genus Colidotea was briefly reviewed by Brusca and Wallerstein (1979b). The 
genus now contains 4 species: C. wstrata (Benedict), southern California; C. findleyi 
Brusca and Wallerstein. Baja California; C. edniondsoni Miller. Hawaii; arfid C. wal- 
Icrsteifii new species. Baja California (Fig. 7). Brusca (in press) discusses the phylogeny, 
evolutionar\ history and zoogeography of Colidotea and its sister-group. Synisonia 
Leach. Both genera are restricted to warm-temperate and subtropical waters of the New 
World (Colidotea) and Old World {Synisoma). Table 1 summarizes the principal fea- 
tures that distinguish these 2 genera from one another and from all other genera of 

A reexamination of the 4 known species of Colidotea warrants an expanded di- 
agnosis of the genus and a key to the species, as follows. 

Diag}iosis. — \doXQ'maQ with body compact, pereon merging smoothly with pleon. 
Cephalon with anterolateral angles produced; with or without a medial cephalic process 
or spine; eyes lateral. Second antennae with flagella comprised of a few. or a dozen or 
so articles, the number increasing somewhat with age (length). Maxillipedal palp of 4 
free articles, apical article large and ovate; epipod present; endite with or without 
coupling hooks. Mandible with large lacinia mobilis. nearly as large, or as large as 
incisor process. Pereon with coxal plates present on segments II-VII; coxae may or 
may not be visible in dorsal aspect, or may be visible only on posteriormost perconites. 
Pleon comprised of single piece, with one pair of lateral incisions (=suture lines); pleon 
length less than '/i total body length; uropods uniramous. Appendix masculinum of 
male arising from base of pleopod 2 endopod. Uropods uniramous. 

Key to the Species of Colidotea 

1. Cephalon with medial spine or process, which may or may not project over 
supra-antennal line 2 

- Cephalon without medial spine or process 3 

2. Cephalic process bifurcate (2-pronged); frontal process subacute (convex); pleo- 
telson with posterior margin angulate; bases of pereopods with dark chromato- 
phore band C edmondsoni 

- Cephalic process simple (single apex); frontal process notched (concave); pleo- 
telson with posterior margin evenly rounded; bases of pereopods without dark 
chromatophore band C. rostrata 

3. Frontal process notched (concave); coxal plates hidden in dorsal aspect; pleo- 
telson posterior margin subacute; eyes large; maxilla 2 trilobate C findleyi 

- Frontal process not notched, but convex; coxal plates visible on pereonites IV- 
VII. in dorsal aspect; pleotelson posterior margin evenly rounded; eyes small; 
maxilla 2 bilobate C wallersteini new species 

Colidotea wallersteini new species 
Figures 4, 5 

7y/7C'^. — Holotype: male, AHF452. Paratypes: AHF4925; SDNHM (1 specimen); 
USNM (1 specimen). 

Locality. — Known from only two localities. Holotype: Mexico. Baja California 
Norte (Pacific coast), Punta Clara, south Rio Santo Tomas, 1 8 December 1 945. collected 
by C. Hubbs (H45-217a). Paratypes: Mexico. Guadalupe Island, off' N coast of Baja 
California. Melpomene Cove, intertidal collections. 18 December 1949. collected by 
R. J. Menzies and D. Reish, I 'elero III (Allan Hancock Foundation), Station No. 1915- 

Diagnosis. — Body straight-sided; coxal plates visible in dorsal aspect on perconites 
IV-VII. Cephalon with supra-antennal line forming 3 points. 1 medial and 2 submedial; 
frontal process distally convex; antenna 1 flagellum a single article; antenna 2 flagcllum 
with about 9 articles; lacinia mobili present on both right and left mandible; molar 



\-/ .;■  . 

FiGi'RF 4. Coliciotca wallcrsietiii new species. H(>loi\pc. AHF 452. Male. 

process simple, greatly reduced (smaller than lacinia); maxilliped with 2 coupling hooks 
on right. 1 on left: maxilla 2 bilobate. smaller lobe with 2-jointed apical process ter- 
minating in single large seta; posterior margin of pleotelson evenly convex. 

/)«c'/7/J//V'//. — Bod\ smooth, straight-sided, with coxal plates on pereonites II-VII 


FiciURE 5. Colidotca wallcrsieini new species. Holotypc. AHF 452. Male. a. right maxilliped. b. maxilla 
c, maxilla 2. d, right mandible, e, uropod. f, pleopod 1. g, pleopod 2. h, pcrcopod IV. i. pcrcopod I. 







rzj 12 

ZJ 11 

Figure 6. Cladistic relationships of the 4 known species of Colidotea and their sister-group. Synisoma. See 
Brusca (in press) for character polarity analysis and discussion of relationships outside this clade. Apomorphies 
indicated are as follow: 1, Reduction from a 5-jointed to a 4-jointed maxillipedal palp. 2. Lacinia mobilis 
of mandible enlarged, subequal to incisor process. 3. Pleon reduced to a single piece, with a pair of lateral 
incisions indicating partial fusion of 1 pleomere. 4, Pleon reduced to a single piece, with no remaining lateral 
incisions. 5. Pleon greatly elongated. '/? or more total body length. 6. Cephalon with an anteromedial process. 
7. Anteromedial process of cephalon bifid. 8. Adoption of a symbiotic relationship with sea urchins of the 
genus Strongylocenlrotiis. and associated morphological adaptations (e.g.. elliptical body; purple pigmenta- 
tion). 9. Eyes enlarged. 10, Posterior margin of pleon acute. 11. Second maxillae with only 2 lobes. 12, 
Typical "spines" of first maxilla outer lobe reduced to 3 slender, stout, simple setae. 

(visible in dorsal aspect only on IV-VII). Length to at least 16.5 mm; length 4-5 times 

Cephalon: Without tubercles or elevations of any kind; broadly immersed in per- 
eonite I; frontal process distally convex. Eyes very small, set on extreme lateral margins. 
Antenna 1 comprised of 4 articles, fourth being the single flagellar article, with terminal 
esthetascs. Antenna 2 with 9-articulate flagellum. Mandible with 4-toothed incisor; 
lacinia with 3 teeth and 3 stout setae; molar process greatly reduced, simple. Maxilla 

1 outer lobe with about 9 small apical setae (some of which arc comb setae) and 3 stout 
simple setae; inner lobe ending in single stout setose spine. Maxilla 2 bilobatc; larger 
lobe with 2 very large plumose setae and about 7 smaller setae, some ctenose; smaller 
lobe with 2-jointed apical process terminating in single large seta. Maxilliped typical 
for genus; enditc with several large apical setae; 2 coupling hooks on right. 1 on left. 

Pereon; Smooth, straight-sided, forming continuous line with pleon; pereonite I 
with anterolateral angles produced to level of eyes, engulhng cephalon. Pereonites 
gradually increasing in width posteriorly; II-VI subequal in length; I and VII somewhat 
shorter than others. Well-developed coxal plates present on II-VII. visible in dorsal 
aspect only on IV-VII. Pcrcopods I-VII slender and ambulatory, terminating in re- 
flexed, biungulate dactyl; sctation simple; ischium of percopod 1 groo\ cd to receive 

Pleon; Posterior margin of pleotelson evenly convex. Uropods simple; with single 
large plumose seta at junction of protopod and lamella. Pleopods typical of genus; 1- 

2 with abundant plumose marginal setae; 3-5 reduced, somewhat fleshy, and cither 



Fici RE 7. Distributions (locality records) of the known species of Colidotea. Open circle = C. wallcislcim. 
Closed circle = C. findleyi. Triangles = C rostrata. C. edmondsoni is endemic to the Hawaiian Islands (not 
on map). 

naked or at most with a few scattered simple setae (as in C. findleyi). Appendix mas- 
culina large, extended beyond apex of pleopodal lamellae, with short apical spines; base 
of pleopod 2 endopod "■muscularized" just below junction of appendix maxulinum. 

Etymology. — Colidotea wallersteini is named in honor of Dr. Barry Wallerstein, 
in acknowledgment of his work on the systematics and ecology of the Idoteidae and 


in recognition of his long-standing friendship: furthermore, the resemblance in form 
is striking. 

Discussion.— Oi' the 4 known species of Colidolea. 2 {C. roslrata and C. edmond- 
soni) share a unique derived character, the large cephalic tubercle. In addition to this 
synapomorphy, they have more general characters in common with one another than 
either shares with C. findlcyi or C. nul/crstcini. No synapomorphies are recognizable 
to unite the latter 2 taxa as sister-species, so their cladistic relationship must be viewed 
as a trichotomy at this time. The character relationships of the known species of 
Colidotea are expressed in the cladogram in Fig. 6. 

Because Hawaii arose as an oceanic island. C. cdniondsoni most likely evolved 
there in isolation subsequent to a dispersal event from western North America. Its 
immediate ancestor was probably a free-living species that was also the parent taxon 
to the commensal C. rostraia. Colidotea findleyi and C. wallersteini can be derived 
from this same ancestral lineage. The latter is strictly a warm-temperate species, known 
only from northwestern Baja California and Guadalupe Island (Fig. 7). Colidotea fin- 
dlcyi is also a warm-temperate species, but exhibits considerable eurythermy in its 
ability to survive the warm summer months typical of the northern Gulf of California. 
The disjunct occurrence o{ C. findleyi on both the western coast of Baja California (and 
Guadalupe Island), as well as the restricted northern Gulf of California population, 
suggests that it may have entered the Gulf during a period of lowered oceanic paleo- 
isotherms (i.e.. Pleistocene glacial periods). Brusca and Wallerstein (1979b) and Wal- 
lerstein and Brusca (1982) have discussed the probability of such events, suggesting 
that the Pleistocene glacials and interglacials (and the associated latitudinal shifts in 
coastal isotherms) resulted in a series of vicariant events such that populations of 
temperate species were repeatedly trapped in the northern Gulf of California as lati- 
tudinal isotherms shifted from north to south and back. These events were probably 
responsible for the evolution of the various northern Gulf endemic species that have 
warm-temperate (Californian) sister-taxa. If this theory is correct, C findleyi can most 
simply be envisioned as a sister-species of C wallersteini, the 2 taxa being vicariant 
products of one of the earlier Pleistocene glacial episodes. Sympatry was later effected 
when a subsequent glacial event facilitated the escape of C findleyi from the Gulf. The 
present interglacial period and relatively high coastal isotherms maintain the current 
disjunct distribution of CJindleyi. 


This study was funded by a grant from the National Science Foundation (DEB 
80-17835). Appreciation is extended to Bill Cooke, George Schultz. and Tom Bowman 
for the loan of specimens of Colidotea cdniondsoni: to the British Museum (Natural 
History) for the loan of Synisoina specimens: and to Fred Schram and Greg Pregill for 
reviewing the manuscript. Frances Runyan executed the two dorsal views. 

Literature Cited 

Brusca. R. C. In press. Phylogeny, evolution and 
biogeography of the marine isopod subfamily 
Idoleinae (Crustacea: Isopoda: Idoteidac). San 
Diego Soc. Nat. Hist. Trans. 

. and B. R. Wallerstein. 1979a. The marine 

isopod crustaceans of the Gulf of California. 
II. Idoteidae. New genus, new species, new rec- 
ords and comments on the morphology, tax- 
onomy and evolution within the family. Proc. 
Biol. Soc. Wash. 92(2):253-271. 

1979b. Zoogeographic patterns of idoteid 

isopods in the northeast Pacihc. with a review 
of shallow-water zoogeography for the region. 
Bull. Biol. Soc. Wash. 3:67-105. 
Iverson, E. W. 1972. A new subtidal Synidolea 

from central California (Crustacea: Isopoda). 
Proc. Biol. Soc. Wash. 85(47):54 1-548. 

Menzies. R. J. 1950. The ta.xonomy. ecology and 
distribution of northern California isopods of 
the genus Idothca with the description of a new 
species. Wasmann .1. Biol. 8(2): 1 55-195. 

, and M. A. Miller. 1972. Systematics and 

zoogeography of the genus Synic/owu (Crus- 
tacea: Isopoda) with an account of Californian 
species. Smithson. Contrib. Zool., No. 102:1- 

Wallerstein. B. R.. and R. C. Brusca. 1982. Fish 
predation: a preliminary study of its role in the 
zoogeography and evolution of shallow-water 
idoteid isopods (Crustacea: Isopoda: Idotei- 
dae). J. Biogeogr. 9:135-150. 


Volume 20 Number 5 pp. 81-94 18 January 1984 

Descriptions of five new muricacean gastropods and comments - - 

on two additional species, in the Families Muricidae LIBRARY 

and Coralliophilidae: (Mollusca) rpf^ "1 4 ',Gp/j 

Anthony D'Attilio and Barbara W. Myers HARVARD 

Department of Marine Invertebrates. San Diego Natural History Museum, San Diego, CA 92112 t/S'/^TY 

Abstract. Five new species of muricacean (superfamily Muricacea) gastropods, one muricid and 
four coraliiophilids, are described as follows: from the Persian Gulf Hexaplex rileyi (Muricidae: Murici- 
nae); from Isla Isabella (= Albemarle Island), Galapagos Islands, Babelomurex deroyorum (Corallio- 
philidae); from Cebu Island, Philippine Islands, Babelomurex jeanneae, Coralliophila armeniaca and 
Coralliophila caroleae (Coralliophilidae). Additional comments and illustrations are provided on pro- 
toconch and opercular characters of Coralliophila neritoidea (Lamarck 1816), the type of Coralliophila. 
and for Coralliophila erosa (Roding 1798). The latter two species occur intertidally and subtidally on 
coral reefs in the tropical Indo-Pacific Ocean. 


Within the Muricidae, the genus Hexaplex Perry, 1811, includes those species with 
a solid and globose shell bearing five to eight spinose more or less foliaceus varices. 
The type species, Hexaplex cichoreum (Gmelin 1791) occurs in the Philippine Islands. 
Specimens from the Persian Gulf, heretofore unknown, are referable to this genus as 
a new species, which we describe in this report. We compare this new taxa to related 
species from the Mediterranean Sea, Indian Ocean and western Atlantic Ocean. 

Four new species of Coralliophilidae are also described here. The Coralliophilidae 
is separable from all other muricacean families in lacking a radula (Thiele 1929, Rob- 
ertson 1970). The members of this family feed suctorially and most are recognized as 
ectoparasites or predators on various species of Cnidaria. These mollusks move from 
host to host, though they are not known to be highly destructive to their prey (Ward 
1 965, Robertson 1 970, 1 980). They have a thaid type operculum with a lateral nucleus. 
Unfortunately shell characters for this family are unstable taxonomic criteria and the 
relationships within the family are complicated further by the lack of a radula. The 
radula is often a distinctive character in the classification of other muricacean taxa at 
the generic level. Twenty-seven genera of Coralliophilidae were enumerated by D'At- 
tilio (1978) and four more have since been added (D'Attilio 1979, Kosuge 1979), 
bringing the total to 31. The two genera considered herein are Coralliophila H. and A. 
Adams, 1853, and Babelomurex Coen, 1922. In Coralliophila a spinose spiral keel is 
wanting and there is little sign of episodic growth. Rather growth takes place in con- 
tinuous increments. Species assigned to Babelomurex possess a spinose shoulder keel 
and may have additional spinose keels on the body whorl. The leading edge of the 
spines usually indicates growth in an episodic manner similar to varices in the Murici- 

In addition to the five new species, two others are discussed: Coralliophila erosa 
(Roding 1798) and C. neritoidea (Lamarck 1816). These two species are widespread 
in the Indo-Pacific. where they inhabit intcrtidal and subtidal coral reefs. Illustrations 
of certain morphological details for these two species have not been published previ- 


Figures 1-4. Hexaplex riteyi D'Attilio & Myers. Dorsal (1) and apertural (2) views of holotype, SDNHM 
81618. Dorsal (3) and apertural (4) views of paratype, SDNHM 81617a. 

The following abbreviations are used: SDNHM (San Diego Natural History Mu- 
seum); AMNH (American Museum of Natural History, New York); and USNM (Na- 
tional Museum of Natural History, Washington, D.C.). 

Systematic Account 

Phylum MoUusca 

Superfamily Muricacea 

Family Muricidae Rafinesque, 1815 

Subfamily Muricinae Rafinesque, 1815 

Genus Hexaplex Perry, 1811 

Type species. — Hexaplex fo/iacea Perry, 1811 by subsequent designation Iredale 
1915 {=Mwex cichoreum Gmelin, 1791). 

Hexaplex rileyi new species 
Figures 1-6 

[Note: Biggs ( 1 973:pl. 5, figs. 8, 9) figured a specimen of Hexap/ex rileyi, but erroneously 
referred it to Murex kUsterianus (Tapparone-Canefri 1875).] 



Figures 5 and 6. Detail drawing of the protoconch of Hexaplex nlcyi from SDNHM 81617b. 

FicH!RES 7 AND 8. Detail drawing of the protoconch of Hexaplex fulvcscens (Sowerby 1834). SDNHM 


Fica RE 9. Detail drawing of the protoconch of Hcxaplex kusterianus (Tapparone-Canefri 1875). SDNHM 


Description.— '^\\e\\ broadly biconic; grayish white with three pale brown bands 
distributed evenly over body whorl, bands mostly distinguishable within outer aperture. 
Protoconch of holotype eroded. Six weakly sloping, tabulate postnuclear whorls; suture 
impressed. Aperture ovate, comparatively large; outer lip crenulate, reflecting external 
sculpture; inner lip white, mostly appressed; anal sulcus well-defined. U-shaped. Si- 
phonal canal open, of moderate length, bent to left and recurved, with five previous 
canal terminations; umbilical chink present. Axial sculpture of nine strong, spinose 
varices on body whorl which cross shoulder and terminate at suture. Five lamellose 
major cords terminate in open spines on each varix; between major cords are numerous 
minor cords of varying width. Entire shell including shoulder finely scabrous. A prom- 
inent single row of open spines on siphonal canal with space separating this row of 
spines from those on body whorl. 


Co/or. — Grayish white with three weak brown bands. 

Tvpe material. -Ho\oXyx)Q: SDNHM 81618. Paratypes: SDNHM 81617a and 
81617b; AMNH 213801; USNM 819632. 

Other material examined. — TweWe specimens from the type locality, Kenneth 
Riley collection. 

Type locality. — 64 km offshore. United Arab Emirates in The Gulf (Persian); depth 
15 m; clinging to petroleum field rigs. 

Dimensions (in mm).— 



















SDNHM 81618 (holotype) 
SDNHM 81617a (paratype) 
SDNHM 81617b (paratype) 
USNM 819632 (paratype) 
AMNH 213801 (paratype) 
12 uncatalogued specimens 
in the K. Riley coll. 

Etymology —^SiTned for Mr. Kenneth Riley, a petroleum engineer who collected 
the specimens. 

Discussion. — The color of this species ranges from pure white to pale brownish 
white with indistinct brown bands. The varices number from six to nine and the 
postnuclear whorls from five to six. Protoconch of paratype SDNHM 81617b has two 
and one-third smooth whorls, somewhat tabulate and of nearly equal diameter. 

This new species is similar to Hexaplex trunculus (Linne 1758) from the Medi- 
terranean Sea. Hexaplex trunculus, however, has a higher spire, is less tabulate, the 
body whorl is more convex and bulges at the shoulder, and the aperture is larger. 

Further comparison can be made with Hexaplex fulvescens (Sowerby 1 834), which 
occurs in the western Atlantic and the Gulf of Mexico, and H. kiisterianus (Tapparone- 
Canefri 1875) from the Indian Ocean. Hexaplex fulvescens has a much larger shell (to 
200 mm), the shoulder is non-tabulate, suture weakly impressed, aperture denticulate, 
and canal much longer. In addition, the varices are concave on their receding side, and 
the shell is white with reddish brown lines over the spiral threads of the entire shell. 
Hexaplex fulvescens has a protoconch of three convex whorls which show very weak 
axial grooves (Figs. 7, 8). Hexaplex kiisterianus has a broad shell with a low spire and 
indistinct suture. The body whorl is very broad and the aperture large. The protoconch 
of H. kiisterianus has an indistinct number of convex whorls (Fig. 9). 

Family Coralliophilidae Chenu, 1859 
Genus Babelomurex Coen, 1922 

Type species. — Fusus babelis Requien, 1848 by original designation. 

Babelomurex deroyorum new species 
Figures 10-14 

Description. S\\e\\ broadly fusiform; spire moderately high; suture weakly im- 
pressed. Protoconch of holotype eroded; teleoconch of six whorls; aperture large, ovate; 
outer lip sharp, with spiral grooves resulting from open spines at margin. Anal sulcus 
broad, shallow; inner lip demarked by elongate node; canal short, open, recurved; 
siphonal fasciole with chink and five short canal terminations. Axial sculpture often 
varices with weakly defined margins on body whorl; penultimate body whorl with nine 
varices; strong shoulder keel separated by gap from two close-set keels below. Narrow, 
flatly triangular, relatively long spines arise at varical margins; spines on keel directly 
below shorter, and those on most anterior keel progressively shorter; presence of ter- 
minal portions of secondary keel obscuring suture; fine squamous threads covering 
entire surface, including upper and lower surface of spines; spiral threads coarser on 


Figures 10-13. Babelomurex deroyorum D'Attilio & Myers. Dorsal (10) and apertural (11) views of ho- 
lotype. SDNHM 81613. Dorsal (12) and apertural (13) views of paratype, SDNHM 81616a. 

Co/or. — White, showing some sHght attrition; small patches are covered with a red 
hydrocoral or bryozoan; paratypes with a lustrous white aperture. 

Type material. - Holotype: SDNHM 81613. Paratypes: SDNHM 8 1 6 1 6a. 8 1 6 1 6b, 
and 8 16 16c; USNM 819633a and 819633b; AMNH 213802a and 213802b. 

Type locality. — Y:>vQdgQd from 75-100 m off Isla Isabella (Albemarle Island). Tagus 
Cove. Galapagos Islands; January 1969. 

Dimensions (in mm).— 

Length Width 

SDNHM 81613 (holotype) 28.8 29.3 

SDNHM 81616a (paratype) 17.0 12.5 

SDNHM 81616b (paratype) 12.2 8.9 

SDNHM 81616c (paratype) 8.4 7.4 

AMNH 213802a (paratype) 13.8 11.0 

AMNH 213802b (paratype) 9.8 9.2 

USNM 819633a (paratype) 12.7 9.0 

USNM 819633b (paratype) 8.0 5.5 

Etymology. — ^dixned for the collectors, Mr. and Mrs. Andre DeRoy. who through 
their collecting efforts, have contributed to our understanding of the endemic molluscan 
fauna of the Galapagos Islands. 


Figure 14. Detail drawing of the protoconch of Bahelonturcx dcroyorum from SDNHM 81616c. 

Discussion. — In addition to the holotype, which is a dead empty shell showing 
some attrition, seven smaller specimens were studied, the largest of which possesses 
five whorls. The keel, showing spines, appears clearly on the third postnuclear whorl. 
Protoconch of SDNHM 81616c has three and one-half whorls with a spiral cord in the 
center and a second spiral cord below; axial threads are diagonal to the protoconch, 
knob-like across the spiral cords. This new species differs from all other eastern Pacific 
and western Atlantic species in the narrow, elongate spines, and finer squamous sculp- 
ture. This species was recovered from the same area as Babc/oniurc.x santacruzensis 
(Emerson and D'Attilio 1970) which has a single row of spines on the shoulder and is 
more closely related to B. dalli (Emerson and D'Attilio 1 963) from the western Atlantic. 
Other eastern Pacific species are B. oldroydi (Oldroyd 1929), B. costata (Blainville 
1832), and B. hindsi (Carpenter 1857). Babeloiuwex oldroydi is found off 
the California coast and has a larger, heavier, coarser shell than B. deroyoruni and 
possesses three spinose keels. Babelonuire.x' costata is similar to B. oldroydi, but with 
less developed and variably keeled spiny cords; B. hindsi is smaller than any of these 

Babeloniurex jeanneae new species 
Figures 1 5-20 

Description. — S\\q\\ biconically fusiform; height from top of aperture to canal ter- 
mination, 1 mm. Protoconch of holotype eroded; six postnuclear angulate whorls with 
large spinose keel at shoulder angle; suture obscured by anteriorly directed, scabrous- 
edged spinose keel corresponding to secondary keel on body whorl; aperture relatively 
large, ovate; columella pillar straight except for slight concavity midway; inner lip edge 
weakly erect anteriorly, outer lip (not entirely mature) extending into the open spines; 
siphonal canal short, broad, open, recurved; fasciole strongly sculptured, with four 
older canal terminations; umbilical chink narrow but deep; seven varices on body 
whorl, eight on penultimate whorl; all varices terminating in lengthy spines. Spiral 
sculpture of a primary row of broad, contiguous, open spines forming the keel; growth 
of spines episodic, giving them a scaly or foliated appearance. Outer portion of primary 
spine bent in direction of growth and entire spiny keel recurved posteriorly; second 
and third row of spines below; bases of secondary spines contiguous and forming keels. 
Five progressively smaller scabrous rows of spines on canal; one similar scaly row 
between second and third row of spines; six rows of weak scabrous spiral cords above 
shoulder terminating at apertural margin. 


Figures 15 and 16. Babclomurex jeanneae D'Attilio & Myers. Dorsal (15) and apertural (16) views of 
hoiotype. SDNHM 79499. 

CoA)/-. — Creamy white, stained with pale ochre in depressed part of shoulder and 
body at receding side of margin. 

'Type material. -Ho\o\\vq: SDNHM 79499. Paratypes: SDNHM 79500 and 81402. 

Other ntaterial exatuined. — OnQ specimen in the Rose D'Attilio collection, one 
specimen in the Barbara W. Myers collection and one specimen in the Donald Pisor 

Type loeality. — BohoX Straits between the Islands of Cebu and Bohol in the Phil- 
ippine Islands. 

DiiJiefisions (in »ini).— 















SDNHM 79499 (hoiotype) 
SDNHM 81402 (paratype) 
SDNHM 79500 (paratype) 
R. D'Attilio coll. 
B. W. Myers coll. 
D. Pisor coll. 

Etyiuology.— Named for Jeanne Pisor. who with her husband Donald Pisor. have 
made noteworthy contributions to the molluscan collections of the San Diego Natural 
History Museum. 

Discussion. — There may be one or two spiny cords below the shoulder keel: the 
number of cords below the secondary keel varies from four to six: growth striae may 
be strong. SDNHM 79500, with a mature outer lip. retains a while intritacalx. a chalky 
white surface layer in some mollusks (D'Attilio and Radwin 1971). with no color other 
than the white surface. The specimen in the R. D'Attilio collection has unusually broad, 
lengthy spines. The protoconch on this specimen has two and one-half whorls: the 
earliest portion is smooth, followed by a sculptured portion with fi\e close-set a.xial 
striae and two spiral cords, beaded where crossed by striae. The specimen in the Barbara 
W. Myers collection is white flushed with a pale violet-pink, and the spines curve 
strongly in the direction of growth. 

Babelomure.x jeanneae probably belongs to a complex of species, the best known 
of which is B. pagodus [of authors, not B. pagodus (A. Adams 1853) (see D'Attilio 
1983)]. Characters similar to B. pagodus are the possession of a spinose keel at the 
shoulder and a secondary keel midway on the body whorl followed anteriorly by a 



Figure 1 7. Detail drawing of the protoconch of Babelomurexjeanneae from specimen m the Rose D'Attiho 


Figures 18 and 19. Babelomurex jeanneae. Specimen from the Rose D'Attilio collection. Detail of spine 

formation showing scales or foliations (18). Detail of broad spines as viewed from above (19). 

FiGLiRE 20. Babelomurex jeanneae. Detail showing spines curved in the direction of growth from specimen 

in the B. W. Myers collection. 

series of progressively diminishing spinose cords. Babelomurex pagodus also has a 
white shell shaded with ochre in the concave portions of the intervarical areas. Varical 
spinose projections also number around eight and are sharply triangulate. Babelomurex 
pagodus is well known and common in southeastern Japan. Based on literature records 
its geographic range is extensive, being known throughout the central and western 
Pacific. As far as is known from the extensive collecting done in Japan, B. jeanneae 
does not occur there. Cebu specimens of B. pagodus resemble in shell morphology 
specimens from southeastern Japan. 

Babelomurex fruiticosus (Kosuge 1979), described from specimens obtained in 
the Straits of Bohoi between Cebu and Bohol Islands, has some similarity to B. jeanneae 
in its possession of a spinose keel at the shoulder and one midway on the body whorl. 
However, the spire is comparatively higher, the area below the shoulder keel shorter, 
appearing compressed, and there are only two spinose cords. The spines are narrow, 
often very elongate, bent or recurved, and their receding side ornamented with sharply 
pointed narrow spinelets imparting to the spines a resemblance to deer antlers. In 
addition, the entire shell is suffused with rich pink or pink-violet, or at times creamy 
white with red-brown at the keel concentrated on the receding side of the varix. 

Babelomurex cristatis (Kosuge 1979) also has conchological characters similar to 
B. jeanneae. However, the eight shoulder keel spines in B. cristatus are shorter, strongly 
up-turned and recurved, with spinelets on their receding side; the secondary keel on 
the mid-area of the body whorl is mostly non-spinose except in mature specimens. 
Two or three inconspicuous cords occur below the mid-cord, and a nearly obsolete 
spiral cord may be present on the canal; the intervarical areas are relatively broad and 
the shell surface is finely and scabrously striate; the color is light to dark tan (flesh), 
occasionally being a deeper brown in the concave areas between the strongly formed, 
rounded, axial costae. 


Figures 21 and 22. Coralliophila armeniaca D'Attilio & Myers. Dorsal (21) and apertural (22) views of 
holotype. SDNHM 79507. 

Genus Coralliophila H. and A. Adams. 1853 

Type species by subsequent designation Iredale 1912: ""Murex neritoideus Chem. 
[nitz]" = Murex neritoideus Gmelin, 1791, not Linne, 1767 = Fusus neritoideus La- 
marck, 1816 (syn. Purpura violacea and P. diversiforniis Kiener, 1836). 

Coralliophila armeniaca new species 
Figures 21-24 

Description. — S\\q\\ fusiform above, compressed anteriorly. Protoconch of holotype 
eroded; six postnuclear convex whorls; suture wavy, distinct; body whorl large, sharply 

Figure 23. Detail drawing of the protoconch of Coralliophila armeniaca from SDNHM 795041. 
Figure 24(a) AND(b). Detail drawing of the operculum o^ Coralliophila armeniaca from holotype. SDNHM 
79507. Internal (a) and external (b) views. 
Figure 25. Detail drawing of the protoconch of Coralliophila rosacea (Smith 1903) from SDNHM 72131. 


FiGL'RES 26 AND 27. Coralliophila cawleae D'Attilio & Myers. Dorsal (26) and apertural (27) views of 
holotype. SDNHM 79505. 

incurved below, flaring out at siphonal fasciole. Aperture large with ten lengthy lirae 
within; lirae at the crenulate outer lip more numerous, reflecting external spiral sculp- 
ture; inner lip erect on lower two-thirds, adherent above. Anal sulcus very weak pos- 
teriorly at juncture of outer lip and columella; siphonal canal open, relatively short and 
broad, recurved, with numerous strong canal terminations on the flaring fasciole, um- 
bilical chink present. Shell with axial sculpture of six swollen, rounded ribs set close 
together, terminating at base of body whorl below and diminishing in strength as they 
abut the whorl above. Spiral sculpture of primary and secondary cords form entire 
surface of the shell; about 15 primary cords with intercalary secondary cords on the 
body whorl below the shoulder; similar cords covering the shoulder; all cords weak to 
strongly scabrous. Operculum thin, translucent, amber-brown with concentric ridges 
externally; internally with two small cords; horseshoe shaped. 

Color. — Deep apricot with slight orange cast; paratypes range from violet to apricot- 
pink. Aperture pale orange at edge of outer lip, white within and on the columella. 

Type material. -\\o\oXypQ: SDNHM 79507. Paratypes: SDNHM 79504a-j; USNM 
819634a and 819634b; AMNH 213803a and 213803b. 

Type locality. — W\ specimens from approximately 75 m depth off^Cebu Island, in 
the Bohol Straits, Philippine Islands; obtained with ground nets. 

Ditnensions (in mm).— 

SDNHM 79507 (holotype) 
SDNHM 79504a (paratype) 
SDNHM 79504b (paratype) 
SDNHM 79504c (paratype) 
SDNHM 79504d (paratype) 
SDNHM 79504c (paratype) 
SDNHM 79504f (paratype) 
SDNHM 79504g (paratype) 
SDNHM 79504h (paratype) 
SDNHM 79504i (paratype) 
SDNHM 79504J (paratype) 
AMNH 213803a (paratype) 
AMNH 213803b (paratype) 
USNM 819634a (paratype) 
USNM 819634b (paratype) 


































Figure 28. Detail drawing of the protoconch of Coral/ioplula caraleac from hoiotype. SDNHM 79505. 
FiciRE 29. Coral/iop/u/a ncniouica (Lamarck 1816). Apertural view of a juvenile SDNHM 66538. 
Pick RF 30. Detail drawing of protoconch of C neritoidea from SDNHM 66538. 

Fkh RE 31 (a) AN[5 (b). Detail drawing of operculum of C. iwriloidca from SDNHM 66538. Internal (a) 
and external (b) views. 

Etymology. — ¥vom arnieniacus, referring to the color of ripe apricot fruit. 

Discussion.— This species appears closely allied to Coralliophila fritschi (von Mar- 
tens 1874) and C. rosacea (Smith 1903), both from South Africa. Another apparently 
closely related species is C. arbutum (Woolacott 1954) { = Rhombothais arhiiiiini Woo- 
lacott 1954) from New South Wales, Australia. These three species differ from C. 
anncniaca by their possession of broadly fusiform shells with a larger body whorl, 
widest at mid-height. In contrast, C. atincniaca has a relatively high spire and the shell 
is broadest well below mid-height. In addition, the angulate shoulder of C. arbimini 
gives it a biconic shape. Protoconch of SDNHM 79504i has three and one-half whorls: 
the first whorl is smooth and rounded whereas the remaining whorls have two spiral 
cords crossed by axial ridges and the nodes are poorly developed where the axial and 
spiral sculpture cross. The protoconch of C. rosacea is illustrated for comparison (Fig. 

Coralliophila caroleae new species 
Figures 26-28 

Description. — 'shcW small, biconic; spire concave, low to moderate height: body 
whorl swollen with moderately angled shoulder tapering to a short, open canal. Pro- 
toconch of nearly three whorls, ridged and beaded. Six postnuclear whorls, moderately 
angled: suture not clear as each succeeding whorl encroaches and somewhat submerges 
previous whorl up to body whorl where suture is clearly defined. Aperture wide, with 
sinuous crenulate outer lip: inner lip smooth, adherent posteriorly: canal short and 
open. Siphonal fasciole composed of a cur\ing scabrous ridge: umbilical chink mod- 
erately deep. Axial sculpture beginning on first postnuclear whorl with nine ribs, in- 
creasing to 14 on subsequent whorls and fading at fourth postnuclear whorl, becoming 


FiGL'RE 32. Coralliophila erosa (Roding 1798). Apertural view of a juvenile SDNHM 77174. 

Figure 33. Detail drawing of the protoconch of Coralliophila erosa from SDNHM 77174. 

Figure 34 (a) and (b). Detail drawing of the operculum of C. erosa from SDNHM 77 1 74. Internal (a) and 

external (b) views. 

barely visible on body whorl. Spiral sculpture of scabrous major and minor cords above 
and below periphery, with strong major cord at periphery of each whorl. Body whorl 
with 1 1 cords between suture and periphery, and 28 cords from periphery to canal; 
width of cords variable; interspaces narrow. 

Color. — Dull orange; aperture orange; inner lip pale orange. 

Type material. -Holoxype: SDNHM 79505. Paratypes: SDNHM 79503 and 81614. 

Other material examined. — One specimen in the Donald Pisor collection and one 
specimen in the James Springsteen collection. 

Type locality. — Boho\ Straits between the Islands of Cebu and Bohol in the Phil- 
ippine Islands, dredged with bottom nets at 75-100 m. 

Dimensions (in mm).- 

SDNHM 79505 (holotype) 
SDNHM 79503 (paratype) 
SDNHM 81614 (paratype) 
D. Pisor coll. 
J. Springsteen coll. 

£'/>'mo/o^. — Named for friend and co-worker, Carole M. Hertz. 

Discussion.— This new species is similar to two other deep water species from 
Mactan Island, Cebu, Philippine Islands: Coralliophila elvirae D'Attilio and Emerson, 
1980, and C. solutistoma Kuroda and Shikama, 1966. Although C soliitistoma was 
described from Japan, it has been recently discovered in the Philippine Islands (D'At- 
tilio and Emerson 1980). 

The spire of C. elvirae is higher and convex rather than concave as in C. caroleae. 
The aperture of C elvirae is restricted to a comparatively longer, narrow opening, with 
the inner and outer lip parallel, whereas C. caroleae has a broad aperture and swollen 
body whorl. The protoconch of C. elvirae consists of two and one-fourth whorls with 
weakly beaded cords; there is only one cord on the final whorl (D'Attilio and Emerson 
1980). The protoconch of C caroleae has two and one-half to three whorls, the beading 
on the cords is much stronger, and the final whorl has two cords. Coralliophila solu- 
tistoDia has a heavier, larger shell with a higher spire and less inflated body whorl; the 
axial ribbing is much stronger and the ribbing is continuous over the entire shell. The 
protoconch is more coarsely beaded (D'Attilio and Emerson 1980). 














Notes on Two Additional Species of Coralliophila 

Two well-known species oi Coralliophila, widely distributed throughout the Indo- 
Pacific, are discussed below. The two species are usually found in such eroded and 
encrusted condition that morphological characters are obscured. Hence, an adequate 
description of the protoconch and early whorls has been lacking in the literature. We 
are fortunate in having at our disposal (SDNHM) juvenile specimens of each, and we 
have thus appended information regarding their shell morphology, protoconch and 
operculum to add to a general understanding of the genus. 

Coralliophila erosa Roding, 1 798 
Figures 32-34 

This species is distributed throughout the Indo-Pacific, mostly intertidal. Mor- 
phologically it exhibits considerable inter- and intrapopulation variation. The shell is 
often, as the name suggests, encrusted or eroded, obscuring the finely scabrous surface. 
It has a characteristic sinuous outer lip, the upper portion of which is deeply concave. 
The operculum is thin with close concentric ridges externally; internally there are six 
U-shaped ridges. The protoconch is distinguished by having one and three-fourths low 
and weakly convex whorls; spire is depressed. An uneroded juvenile specimen (SDNHM 
77174) from Hawaii is illustrated. 

Coralliophila neritoidea Lamarck, 1816 
Figures 29-31 

This species has a distribution and habitat similar to Coralliophila erosa. The white 
shell is most often encrusted with calcareous organisms. When not encrusted or eroded, 
the surface of the shell is spirally scabrous. After the first two to three postnuclear 
whorls there often follows a rapid expansion of the body whorl. The purple aperture 
is less prominent in immature specimens. The operculum is a dark chocolate brown 
with a lateral mid-central nucleus. Internally it has a thickened ridge and four ovate 
ridges spaced over the remaining surface. The protoconch has two to two and one-half 
smooth, acutely conical whorls, followed by a whorl with a midway rib. A juvenile 
specimen (SDNHM 66538) is illustrated. 


The following friends and collectors have contributed towards the completion of 
this paper: Mr. Kenneth Riley, for donating the specimens used in the description of 
Hexaplexrileyi; Mr. and Mrs. Andre DeRoy from Santa Cruz Island, Galapagos Islands, 
for donating specimens of Babelomurex deroyorunr, Mr. Donald Pisor of San Diego, 
California, and Mr. Loyal J. Bibbey of Imperial Beach, California, for donating spec- 
imens of Babelomurex Jeanneae: Mr. Gene Everson of Lauderhill. Florida, and Mr. 
Donald Pisor for contributing specimens of Coralliophila armeniaca: Mr. Donald Pisor 
and Mr. Jim Springsteen of Manila, Philippine Islands, for donating specimens of 
Coralliophila caroleae. 

We are especially grateful to Mr. David K. MuUiner of San Diego, California, for 
the photographs used in this paper. We further thank Dr. Richard C. Brusca, Chairman, 
Department of Marine Invertebrates, San Diego Natural History Museum; Dr. William 
K. Emerson, American Museum of Natural History; and Dr. Emily H. Yokes, Tulane 
University, for editorial review of the manuscript. Marjorie Rea kindly typed the 

Literature Cited 

Adams, A. 1853. Descriptions of several new Biggs. J. E. J. 1973. The marine mollusca of the 

species of Murex. Rissoina. Planaxis and Eu- Trucial coast. Bull. Br. Mus. (Nat. Hist.) Zool. 

lima from the Cumingian collection. Proc. Zool. London 24(8):343-42 1 . 

Soc. London for 1851 19:267-272. BlainviUe. H. M. 1832. Disposition methodique 

Adams, H. and A. 1853-54. Genera of Recent des especes recents et fossiles des genres 

mollusca. Van Voorst, London, vol. 1. 484 pp. Pourpre. Ricinule. Licorne et Concholepus de 


M. de Lamarck, et description des especcs nou- 
velles ou peu connucs, faisant partic de la col- 
lection du Museum d'Histoire Naturcile de 
Paris. Nouv. Ann. Mus. d'Hist. Nat.. Paris, 1: 
Carpenter, P. P. 1857. Report on the present state 
of our knowledge with regard to the Mollusca 
of the west coast of North America. Rep. Brit. 
Assoc. Adv. Sci. for 1856, pp. 159-368. 
Chenu. J. C. 1859. Manuel de conchyliologie et 
de paleontologie conchyliologique. Paris, vol. 
1,508 pp. 
Coen, G. 1922. Del genere Pseudomurex (Mon- 
terosato, 1872). Atti. Soc. Ital. Sci. Nat. Mus. 
Civico. Storia Nat. Milano 61:68-71, pi. 2. 
D'Attilio, A. 1978. Catalogue of the Family Cor- 
alliophilidae. Festivus 10(10):69-96. 

. 1979. Corrections and additions to the 

Catalogue of the Family Coralliophilidae. Fes- 
tivus ll(10):83-85. 

1983. Concerning the identity of Murex 

pagodus A. Adams, 1853. Festivus 15(1): 

D'Attilio, A., and W. K. Emerson. 1980. Two new 
Indo-Pacific Coralliophilid species (Gastro- 
poda, Muricacea). Bull. Inst. Malac. Tokyo 1(5): 

D'Attilio, A., and G. Radwm. 1973. The intri- 
tacalx, an undescribed shell layer in moUusks. 
Veliger 1 3(4):344-347. 

Emerson, W. K., and A. D'Attilio. 1963. A new 
species of Latiaxis from the western Atlantic 
(Mollusca, Gastropoda). Amer. Mus. Novi- 
tates 2149:1-9. 

, and . 1970. Three new species of 

Muricacean gastropods from the eastern Pa- 
cific. Veliger 12(3):270-274. 

Gmelin, J. F. 1791. Caroli a Linne Systema na- 
turae per regna tria naturae. Ed. 13. Leipzig. 
Germany, vol. 1, pt. 6, pp. 3021-3910. 

Iredale, T. 1912. New generic names and new 
species of marine Mollusca. Proc. Malac. Soc. 
London for the year 1914 10:217-228. 

. 1915. A commentary on Suter's "Manual 

of New Zealand Mollusca." Trans, and Proc. 
Royal Soc. of New Zealand 47:417-497. Wel- 

Kiener, L. C. 1836. Species general et inconogra- 
phie des coquilles vivantes comprenant la col- 
lection du Museum d'Histoire Naturelle de 
Paris .... Genre Pourpre {Purpura. Lam.), pp. 

Kosuge, S. 1979. Descriptions two new subgenus 
[sic] and seven new species of the genus La- 
tiaxis (Gastropoda, Mollusca). Bull. Inst. Ma- 
lac. Tokyo l(l):3-8. 

Kuroda. T.. and T. Shikama, in T. Shikama. 1 966. 
On some new Latiaxis and Coral/iophila in 
Japan. Venus 25(l):21-25. 

Lamarck. J. B. P. A. de M. de. 1816. Tableau 
encyclopedique et methodique des^trois regnes 
de la nature. Liste. 16 pp. 

Linne, C. von. 1758. Systema naturae. Stock- 
holm. 10th ed. vol. 1. pp. 1-824. 

. 1767. Systema naturae. Stockholm. 12th 

ed. Vol. I Pt. II-Mollusca. pp. 1081-1267. 

Martens, E. von. 1874. Uebereinige Sudafrikan- 
ische mollusken. Jahrb. Deutsch Malak, Ge- 
sell. pp. 119-146. 

Oldroyd, L S. 1929. Description of a new Cor- 
alliophila. Nautilus 42(3):98-99. 

Perry, G. 1811. Conchology or the natural history 
of shells, containmg a new arrangement of the 
genera and species. Wm. Miller, London. 

Rafinesque, C. S. 1815. Analyse de la nature ou 
tableau du univers et des corps organises. Bar- 
ravecchia, Palermo. 

Requien, E, 1848. Catalogue des coquilles de I'lle 
de Corse. Avignon. 

Robertson, R. 1970. Review of the predators and 
parasites of stony corals with special reference 
to symbiotic prosobranch gastropods. Pac. Sci. 

. 1980. Epitonium nullecostatuni and Cor- 

alliophila clathrata: Two Prosobranch gastro- 
pods symbiotic with Indo-Pacific Palythoa 
(Coelenterata: Zoanthidae). Pac. Sci. 34(1):1- 

Roding, P. F. 1798. Museum Boltenianum. Ham- 
burg. 199 pp. 

Smith, E. A. 1903. A list of species of mollusca 
from South Africa, forming an appendix to G. 
B. Sowerby's "Marine Shells of South Africa." 
Proc. Malac. Soc. London 5:354-402. 

Sowerby, G. B. II. 1834-41. The Conchological 
Illustrations, Murex. Sowerby, London, pis. 58- 
67. 1834; pis. 187-199, 1841. 

Tapparone-Canefri, C. M. 1875. Viaggio dei . . . 
O. Antinori, O. Beccari ed. A. Issel nel Mar. 
Rosso . . . 1870-71. Studio monografico sopra 
il Muricidi, etc. Ann. Mus. Civ. Stor. Nat. Gen- 
ova 7:560-640. 

Thiele, J. 1929-35. Handbuch der Systemati- 
schen Weichtierkunde, Jena: Gustav Fischer, 
Bd. I, Tiel I, 376 pp. 

Ward, J. 1965. The digestive tract and its relation 
to feeding habits in the stenoglossan proso- 
branch Coralliophila ahbrexiata (Lamarck, 
1816). Canadian Jour. Zool. 43:447-464. 

Woolacott, L. 1954. New shells from New South 
Wales. Proc. Roy. Soc. New South Wales. Zo- 
ology 1952-53: pp. 37-39. 






Volume 20 Number 6 pp. 95-98 18 January 1984 

The Fossil Leptostracan Rhabdouraea bentzi (Malzahn, 1958) 

Frederick R. Schram 

Department of Geology; Paleontology Section. San Diego Natural History Museum, San Diego. CA 
92112 USA 

Eric Malzahn 

Aufder Heide 33. D-3004 Isernhagen, German Federal Republic 

Abstract. The type material of the only known fossil leptostracan phyllocarid, Rhabdouraea bentzi, 
is reexamined. This species was originally placed in the genus Nebalia. The distinctive caudal rami and 
the form of the carapace, however, require separate generic and familial status for this Permian material. 
A redefinition of the living family Nebaliidae is also provided. 


The fossil record of the Phyllocarida has always presented problems for phylo- 
genetic analysis because of the incomplete nature, generally, of the fossils (Rolfe 1981). 
This has been especially vexing for the assessment of the living order Leptostraca where, 
with a single exception, fossils are non-existent. Malzahn (1958) described Nebalia 
bentzi on the basis of a single specimen from the Upper Permian Zechstein of West 
Germany. Further discussion of this species was provided by Glaessner and Malzahn 
(1962). Initial assignment of this material to the living genus Nebalia reflected a con- 
servative approach pending the discovery and study of more and better material. 
Unfortunately, despite continuing efforts at collection and study of this fauna by the 
junior author, no additional specimens have turned up. However, restudy of the known 
material has enabled us to clarify the relationship of this Permian species to the living 

Systematic Paleontology 

Class Malacostraca Latreille, 1806 

Subclass Phyllocarida Packard, 1879 

Order Leptostraca Claus, 1880 

Family Rhabdouraeidae n. fam. 

Diagnosis. — S>?LmQ as that of the genus. 
Type genus.— Rhabdouraea n. gen. 

Rhabdouraea new genus 

D/fl^/7m«.— Carapace short, not covering abdomen. Caudal rami rod-like, at least 
as long as abdomen. 

Etymology. — Yrom. the Greek rhabdos (=rod) and uraea (=tail), gender feminine. 
Type species.— Nebalia bentzi Malzahn, 1958. 

Rhabdouraea bentzi (Malzahn. 1958) 

Diagnosis.— Since only one species is known, the diagnosis of the species is the 
same as that of the genus. 



Figure. 1 . Partial reconstruction of Rhahdouraea bentzi (Malzahn). A.) lateral view of abdomen. B.) dorsal 
view of telson and caudal rami. Scale 2 mm. 

Holotype.—Z45, in the collection of the Niedersachsisches Geologisches Lande- 
samt, Hannover. From a depth of 280-282 m. Shaft 4, of the Friedrich-Heinrich Mine, 
near Ort Hoerstgen, Niederrhein, German Federal Republic. Zechstein 1, Upper Perm- 

Description.— The posterior margin of the carapace is visible on the type, and 
covers the posterior aspect of the thorax while leaving the anterior pleomeres exposed 
(Figs. 1 and 2). The pleomeres are subequal, each about 0.7 mm in length. The posterior 
margin of the abdominal tergites are raised as a slight ridge. The abdominal pleura are 
rounded anteriorly and somewhat acuminate posteriorly. At least the four anterior 
pleopod protopods are long and robust; the fifth and sixth are present, appear to be 
somewhat smaller than those of the anterior limbs, but cannot be clearly discerned 
because of the preservation of the specimen. The seventh pleomere appears to lack 
appendages. The distal branches of the anterior pleopods are robust, and marked by a 
line of lateral pits (which may have been sockets of marginal setae). The telson is only 
slightly shorter than the pleomeres, and with a slight fossa or depression on the dorsal 
surface between the bases of the caudal rami. The telson bears terminally a set of large, 
rod-like, papillose caudal rami. The length of these rami cannot be determined exactly 
since their distal ends are broken off. However, the longer ramus (now broken in two 
pieces) is at least 3.5 mm and indicates the rami were probably at least as long as the 

Family Nebaliidae Baird, 1850 

Diagnosis.— Carapace generally large, covering anterior pleomeres as well as the 
thorax. Caudal rami relatively short, as flaps with marginal setae. 
Type genus.— Neba/ia Leach, 1814. 


The two distinct features of Rhahdouraea bentzi are the short carapace and the 
long, rod-like caudal rami, so diagnostic in fact as to warrant separate generic and 
familial status for this species. 


FiGiRF 2. Rhabdouraea bentzi (Malzahn). holotype. Z45. A.) Right side [right caudal ramus visible on 
original illustrations since broken off. but still retained with specimen], lOx. B.) Left side, closeup of anterior 
pleopods and caudal ramus, 16x; c— carapace, cr— caudal ramus, pi — pleuron of second pleomere, pr— 
pleopodal protopods. r— distal rami of pleopods. t — telson. 

The living leptostracans typically have a large carapace that completely encloses 
the body, except for the posteriormost portions of the abdomen, and lobate. setose 
caudal rami. The only exception to these features occurs in the genus Nehaliopsis. This 
pelagic form has the posterolateral aspect of the carapace truncate, exposing the pleo- 
meres and ventral portions of the posterior thoracomeres. In addition, the caudal rami 
in Nebaliopsis are thin and leaf-like. These features are prompting Hessler (in prep.) 


to place Nehaliopsis in its own family, separate from the benthic nebaliid genera. The 
differences noted on Rhahdouraea, incomplete though the fossil is, are of a magnitude 
at least as great as that which separates Nehaliopsis from Nehalia. Paranebalia, and 
Neba/ic/la, and thus justify separating this Permian taxon by itself 

All the living nebaliids, benthic or pelagic, have relatively short or modest sized 
caudal rami with setose margins. These rami function in a manner analogous to uropods, 
and assist the animal in swimming. The very long, papillose, rod-like rami in Rhah- 
douraea are unlike any of those seen among leptostracans, and are more akin to those 
seen in notostracan branchiopods. 

Several features of Rhahdouraea hentzi ally this species to the leptostracan phyl- 
locarids, viz., the four well-developed anterior pleopods, the two apparently small 
posterior pleopods, the lack of the seventh pleopods, the robust pleopod protopods, 
the lateral row of pits on the pleopod distal branches, and the telson not developed 
dorsally between the caudal rami. The recognition of the separate familial status of 
Rhahdouraea hentzi seconds the suspicions of Rolfe (1969) that this species was not 
referable to any of the Recent genera. This requires an emendation of the diagnosis of 
the living family Nebaliidae noted above. 


Drs. Robert Hessler and Ian Rolfe examined the material and offered valuable 
counsel on the taxonomic issues. Work was supported in part by NSF grant DEB 79- 
03602 (FRS). 

Literature Cited 

Malzahn, E. 1958. Eine neuer jungpalaozoischer Rolfe, W. D. I. 1969. Phyllocarida. Pp. R296- 

Krebs aus dem niederrheinischen Zechstein. R331 in R. C. Moore (ed.). Treatise on Inver- 

Zeit. deutsch. geol. Ges. 1 10:352-359. tebrate Paleontology, Part R, Arthropoda 4(1), 

Glaessner, M. F., and E. Malzahn. 1962. Neue Geol. Soc. Am. and Univ. Kansas Press, Law- 

Crustaceen aus dem niederrheinischen Zech- rence. 

stein. Fortschr. Geol. Rheinld. u. Westf. 6:245- . 1981. Phyllocarida and the origin of the 

264. Malacostraca. Geobios 14:17-27. 




UNlVERBiTY Volume 20 Number 7 pp. 99-134 18 January 1984 

Phylogeny, evolution and biogeography of the marine isopod 
Subfamily Idoteinae (Crustacea: Isopoda: Idoteidae) 

Richard C. Brusca 

Department of Marine Invertebrates, San Diego Natural History Museum. San Diego, California 92112 USA 

Abstract. The patterns of spatial distribution attained by the genera of Idoteinae are discussed in 
hght of a cladistic analysis of the suborder Valvifera and the subfamily Idoteinae. A schematic pattern 
analysis technique is demonstrated and reveals the probability of multiple origins of similar pleonal 
morphologies among various genera of Idoteinae. Reduction in the maxillipedal palp has occurred 
numerous times within the Idoteinae. while loss of the biramous uropodal condition has probably 
occurred twice. A geographic cladogram of temperate Gondwanan shores is proposed. An evolutionary- 
biogeographic narrative is presented, in which a set of hypotheses is developed to describe the history 
of the Idoteinae in time and space. The subfamily Idoteinae appears to form two principal lines of 
descent, both arising in the Triassic or Jurassic. One of these lines remained closely tied to the Southern 
Hemisphere (primarily Old World) temperate marine shores from which the Idoteinae is derived. The 
other line invaded the Northern Hemisphere and various New World environments, and more recently 
(Cenozoic) underwent a radiation in the American tropics. The success of this latter lineage (e.g., 
Erichsonella. Eusymmerus, Parasymmerus. Cleantioides) may be due to certain morphological and life 
history adaptations not found in New World species of the former line (e.g., Idotea. Synidotea). The 
Valvifera probably originated in the temperate Southern Hemisphere, at least by Permean/Triassic 
times. Global distribution patterns of some genera can be ascribed most parsimoniously to vicariance 
processes, and in others to dispersal, ecological phenomena, or a combination of processes. Other factors 
have apparently also affected modern distributional patterns of idoteine genera, for example, extinctions. 
Biogeographical data can be used to elucidate viable alternative cladistic hypotheses to those generated 
solely on parsimonious patterns of synapomorphy. Biogeographic data can also be used, in conjunction 
with the cladogram, to identify probable ancestral taxa. 


Few groups of marine invertebrates have enjoyed analysis by cladistic techniques, 
despite the current popularity of the method. The only previous attempt to examine 
the phylogenetic relationships of an isopod taxon by strict cladistic (Hennigian) meth- 
odology was that of Williams ( 1 970), who analyzed the relationships of North American 
epigean species oi Asellus (Asellota): but, as was common with early attempts at Hen- 
nigian analyses, he used a weakly defined method of character state polarity assessment. 
Despite this, he was still able to construct a very plausible ph\ logenetic hypothesis, or 
cladogram. for the 14 species he treated. 

One reason for the paucity of cladistic studies on marine invertebrates is the 
necessity to work with a group whose taxonomic relationships arc reasonably well 
known within the context of the larger hierarchical taxon to which the study group 
belongs. Without this knowledge, an assessment of character polarity is difficult to 
obtain, and without polarities, construction of both cladograms and phylogenetic (evo- 
lutionary) trees must- be based on speculatively generated hypothetical ancestors (ex- 
pressed or implied). Although several marine invertebrate taxa are well understood 
systematically (e.g.. certain families of molluscs, crabs, barnacles), the great majority 
are not. Several crustacean isopod groups are also well understood in this regard. The 
subfamily Idoteinae (suborder Valvifera; family Idoteidae) is one such group. 

The present study attempts to answer questions about the evolutionary history of 
the Idoteinae using cladistic techniques. Specific questions addressed arc: What are the 


phylogenctic patterns and relationships of the genera of Idoteinae? What are the spatial 
patterns and relationships of the genera of Idoteinae? What evolutionary history is 
suggested when these patterns are compared to one another and to the earth's geological 
and ecological history? Previous studies on these topics include comments on the 
phylogeny and biogeography of the genus Idotea by Menzies (1950c/): on the genus 
Synidotea by Menzies and Miller (1972); and on the comparative morphology of the 
valviferan higher taxa in general by Sheppard (1957). A review of the distribution of 
shallow-water idoteine species in the northeastern Pacific was given by Brusca and 
Wallerstein (1979/)). and a discussion of the possible ecological and historical mech- 
anisms regulating distribution and latitudinal trends in morphology and behavior in 
that taxon is provided by Wallerstein and Brusca (1982). 


The methods used in this study are largely summarized in 3 recently published 
books (Eldredge and Cracraft 1980, Nelson and Platnick 1981, Wiley 1981). However, 
even the principal spokesmen of current cladistic theory are not without disagreement 
on both details of procedure and certain underlying philosophical issues. As Eldredge 
and Cracraft (1979) point out, "No two cladists agree with each other (or, for that 
matter, with Willi Hennig) on every point, and this 'school' of systematics is no more 
a monolith than that of the more traditional 'evolutionary taxonomy'."' The overall 
concept of cladistic or "phylogenctic" analysis has evolved considerably since Hennig 
(1966). and indications are that it will continue to change for some time to come. For 
these reasons, and others. I offer the following position statements. 

While the present study is cladistic in nature, it is my opinion that such analyses 
are most useful as investigatory techniques and do not represent the final word on 
phylogeny. The most powerful (and important) aspect of cladistic methodology is its 
ability to posit and define monophyletic groups in an unambiguous and testable manner. 
Synapomorphy patterns, however, do not constitute the sole source of phyletic infor- 
mation on a taxon. but rather must be compared to other kinds of data and analyses 
when constructing phylogenctic trees, evolutionary scenarios, AND classifications. A 
cladogram depicts only a sequence of character appearances, which may or may not 
correspond to speciation events (Hull 1979). There appear to be 4 principal products 
that can result from phylogenctic analyses: cladograms, phylograms (evolutionary trees), 
evolutionary scenarios, and classifications. The cladogram should be viewed as a "best 
guess" in the face of uncertainty (Felsenstein 1973, Harper 1979, Hull 1979, Simberloff 
et al. 1981, Endler 1982), and information contained in any of these other products of 
phylogenctic analyses can legitimately be used to improve any other, including the 
cladogram itself (Hull 1979). In the present study I construct a cladogram, a phylogram, 
and an evolutionary scenario for the subfamily Idoteinae. and use the latter two products 
to shed new light on, and make ammendments to, the cladogram. 

Nomenclature and general terminology are taken from current literature on val- 
viferan isopods (see above references). Morphological structures discussed in this paper 
are illustrated in Fig. 1. Analysis of character polarity is based on out-group analysis 
(see Eldredge and Cracraft 1980. de Jong 1980, and Watrous and Wheeler 1981). I 
believe that out-group comparisons need not be rigidly constrained by nomenclatural 
rank or Linnean hierarchical structure, but are applicable at all levels of a cladogram. 
Though parsimony is a potent methodological tool, it is primarily a method of logical 
analysis, not a biological law or principle. Application of parsimony should be an initial 
technique, or one to be used in the absence of other data. To continue to hold to the 

FiciURF 1. Aspects of the morphology of idoteid isopods discussed in the text, a, Synidotea harfordi; note 
multiarticulate fiagclla on antennae 2 and 0+1 pleonal morphology, b, Cleantioides occidentalism note un- 
iarticuiate (clavate) flagcllae on antennae 2 and 3+1 pleonal morphology, c, Colidotca findleyi: note mul- 
tiarticulate fiagcllae on antennae 2 and 0+1 pleonal morphology, d. Mandible of Kusyninwrus antennatus; 


note large 4-toothed incisor, smaller lacinia mobilis. and stout molar process, e. Mandible of Colidotea 
findleyi; note 5-toothed incisor, large lacmia mobilis. and stout molar process, f, Maxilliped of Colidotea 
findleyi\ note 4-articulate palp, g, Uniramous uropod of Colidotea findleyi. h. Antenna 2 of Colidotea findleyi; 
note multiarticulate flagellum. i. Antenna 1 of Colidotea findleyi; note uniarticulate flagellum. j. Antenna 2 
of Erichsonella cortezi: note uniarticulate (clavate) flagellum. k, Plcopod 2 of Enchsonella cortezi (male); 
note appendix masculinum. 


"simplest" explanation (i.e.. the shortest cladogram) in the face of biological evidence 
indicating a less parsimonious but more biologically reasonable explanation is both 
nonscientific and an abuse of the tool. There is little point in creating a falsifiable 
hypothesis if one docs not accept all forms of data that can falsify it. In this regard I 
agree with Kitts (1981) that phylogenetic patterns (and hence analyses) ARE*historical 
in nature and this involves describing the real world; one could not be writing history 
if one supposed every relationship between events to have transpired in the "most 
direct" manner. Whether or not any particular phylogeny is parsimonious is something 
to be found out in the course of a historical investigation, it is not something to be 

Finally, my technique for the historical biogeographic analysis follows the hypo- 
thetico-deductive method, primarily as described by Morse and White ( 1979). I assume 
no particular paradigm to be of overriding importance, but rather attempt to interpret 
the patterns of characters and distributions in the most parsimonious (biological par- 
simony) fashion possible. McDowall (1978) was. of course, correct in noting that one 
can never know with certainty whether any given individual component track is the 
product of vicariance or dispersal. However, that both phenomena exist in nature can 
hardly be denied, and it has been my task in the present study to decide, where possible, 
which of these two phenomena (or others) produced the biogeographic patterns seen 
today in the genera of Idoteinae. Needless to say. I have been severely hampered in 
this regard by the absence of cladograms for other coastal marine taxa. 

Higher Classification of the Order 


Out-group analysis requires acceptance of some higher level classificatory structure 
before statements regarding the relationships of lower, inclusive taxa can be made. For 
the purposes of this study, I accept the monophyletic nature {scnsii Hennig, 1966) of 
three taxa: the order Tanaidacea, the order Isopoda, and the suborder Valvifera. The 
monophyletic nature of these distinct taxa are, to my knowledge, unquestioned. 

While the nature of the primitive isopod body plan (presented below) is generally 
agreed upon, the relationships of the 9 isopod suborders are unknown and fraught with 
speculation (see Schultz 1979 for recent summary comments). Various authors have 
described the nature of the primitive or ancestral isopod morphotype, which is char- 
acterized by the following combination of characters: carapace wanting; pereopods 
uniramous; respiratory structures (heart and branchial surfaces) primarily abdominal; 
pereopodal coxae forming marginal plates on pereonites; first and second antennae 
with multiarticulate flagella; mandible with a multiarticulate palp; appendix masculina 
present only on second pair of pleopods; uropods biramous (probably attached ter- 
minally or subterminally to telson or pleotelson, although some authors suggest a lateral 
attachment vis-a-vis the cirolanoids); eyes entirely sessile; all pereopods more or less 
similar; pereon of 7 free somites (thoracomeres 2-8); pleon of 6 free somites and a 
telson (or possibly 5 free somites and a pleotelson); maxilliped with a large basal endite 
and reduced endopodal articles (the latter forming the 5-articulate palp); maxilliped 
with a small, ovate, nonbranchial epipodite (the "endognath"); penes and opening of 
vas deferens on thoracomere 8; simple foregut; and maxillary glands present in adults. 

This generalized ancestral isopod plan was first developed in the early studies of 
Bate and Westwood (1861-1868), Stebbing (1893), and Caiman (1909), and more 
recently by Schram (1974) and Hessler et al. (1979). The concept of this morphotype 
is supported by fossil data as well as by comparison with other peracarid and mala- 
costracan taxa. It is also compatible with all three "competing" hypotheses of extant 
primitive isopod morphotypes (i.e., cirolanoid, phreatoicid, asellote). 

The Valvifera stand apart as perhaps the most distinct of the isopod suborders in 
several regards. Important features distinguishing the valviferan body plan are (see Fig. 
1): (a) coxae of thoracic legs (pereopods) with both dorsal and ventral plates, the latter 
extending over the sterna; (b) uropods attached laterally on pleotelson. but modified 


to form ventral opercular plates covering the pleopods; (c) vas deferens (and penes) 
opening on abdomen of male, rather than on thorax as in all other isopods, excepting 
the Oniscoidea (i.e., on pleonite 1 or on the articulation of pleonite 1 and thoracomere 
8); (d) flagella of first antennae reduced to one or a few vestigial articles; (e) pleon of 4 
or fewer free somites (plus the pleotelson); (0 uropods biramous or uniramous; (g) 
maxillipedal palp of 3-5 articles: (h) second antennae uniramous. fiagellum multiar- 
ticulate or uniarticulate; (i) mandible with or without a 3-jointed palp. Attributes (a) 
and (b) are unique synapomorphies that distinguish the Valvifera from all other isopod 
taxa; attributes (c) through (0 are valviferan synapomorphies that also appear in one 
or more other isopod suborders (apparent convergences). 

The current classification of the isopod suborder Valvifera is as follows: 

Order Isopoda Latreille. 1817 
Suborder Valvifera Sars, 1882 

Family Holognathidae Thomson, 1904 
Family Idoteidae Fabricius, 1798 

Subfamily Idoteinae Dana, 1852 

Subfamily Parachiridoteinae Elkaim and Daguerre de Hureaux, 1976 

Subfamily Glyptonotinae Miers, 1881 

Subfamily Chaetilinae Dana, 1852 (=Macrochiridoteinae Nordenstam. 1933) 

Subfamily Mesidoteinae Racovitza and Sevastos, 1910 
Family Xenarcturidae Sheppard, 1957 
Family Arcturidae G. W. Sars, 1897 
Family Amesopodidae Stebbing, 1905 
Family Pseudidotheidae Ohlin. 1901 

The relationships of the 6 valviferan families have long been unclear. The only 
cogent discussion of the topic was that of Sheppard (1957). The cladogram in Fig. 2 
depicts the best arrangement that I have been able to devise for these families, being 
the most parsimonious, and admitting no convergences, parallelisms or reversals. A 
convincing higher level classificatory scheme of the 9 isopod suborders does not pres- 
ently exist, and carcinologists disagree over the relationships among these taxa. For 
this reason, the Tanaidacea was used as an out-group to construct the cladogram of 
valviferan families (Fig. 2). Tanaidacea is the peracarid order "traditionally" (Schram 
1981) taken to be the most probable sister-group to the Isopoda (also see Slewing 1963 
and Fryer 1964). Whether or not it is the actual sister-group of the isopods is unim- 
portant for its use in out-group comparison, however, as it is clearly a closely related 
taxon within the unified peracarid line. Character polarity assessments based on tanaids 
were compared to those obtainable by using the Amphipoda. Cumacea and hypothetical 
ancestral isopod as out-groups and no changes in polarity were required when these 
other groups were used in place of tanaids.' A step-by-step discussion of the cladogram 
of valviferan families follows, the numbering in the text following that of Fig. 2. 

Tanaids are united to the isopods only by possession of their peracarid attributes, 
the most obvious of these being: (a) pereopodal coxae with thin ventral plates (oos- 
tegites) that form a female brood pouch for the developing young: (b) mandibles with 
lacinia mobili in adult stages of life cycle, and (c) young released from the marsupium 
in subadult "mancoid" stage. At this level of analysis these attributes are symplesio- 
morphies: I know of no synapomorphies unique to the tanaids and isopods. 

The Isopoda are united by the features listed above for the primitive isopod 
morphotype. The first 4 of these are synapomorphies, as follow: (1) carapace wanting 
(vs. present in tanaids): (2) pereopods uniramous (vs. retaining vestiges of exopods); 
(3) respiratory structures (branchial pleopods and heart) primarily abdominal (vs. tho- 
racic): (4) pereopodal coxae forming marginal plates on pereonites (vs. not forming 
plates). - 

The Holognathidae shares in common with its sister-group (the remaining 5 val- 
viferan families) the 4 valviferan synapomorphies listed earlier: (5) pereopodal coxae 
form ventral (sternal) plates; (6) uropods modified into opercular plates covering pleo- 



























































Figure 2. Cladogram of families of Valvifera. Closed boxes indicate apomorphies; open boxes plesio- 
morphies. Numbering of characters corresponds to text discussion. 

pods; (7) vas deferens and penes opening on abdomen of male, rather than on thorax; 
(8) flagella of first antennae reduced to one or a few vestigial articles. The Holognathidae 
is a monotypic family containing but one species, H. stewarti (Filhol). It cannot be 
distinguished by an autapomorphy of its own, although it stands apart from all other 
species in the suborder Valvifera by its retention of a palp on the mandible, as well as 
numerous other primitive attributes (e.g., biramous uropods, 5-articulate maxillipedal 
palp, pleon of 4 free somites plus the pleotelson). Holognathus may be considered a 
relict taxon within the Valvifera. Nordenstam (1933) long ago recognized the lack of 
apomorphies in Holognathus, suggesting that it might best be incorporated into the 
Idoteidae. Were this done, however, Holognathus would probably have to be ranked 
as a sixth subfamily, rather than included in the Idoteinae as Nordenstam suggested. 
The presence of 4 free pleomeres places this genus at a position ancestral to both the 
Idoteinae and the Glyptonotine-group discussed below (see Fig. 3). The remaining 
valviferan families are thus united by the loss of the mandibular palp (9). Clearly, were 
one to place Holognathus in the Idoteidae, characters 5-8 would become synapomor- 
phies uniting Idoteidae to all other valviferan families. 

The Idoteidae stand apart as the only family in which the uropods may be reduced 
from the primitive biramous state to a uniramous condition (10), and in which a 
reduction of the maxillipedal palp takes place (11). The Idoteidae have been charac- 
terized by two other "synapomorphic trends": trends towards fusion of the pleonites 
and towards fusion (or loss) of the flagellar articles on the second antennae. These 
reductions, however, are convergent to patterns that also occur in other valviferan 
families and thus have not been used to construct the cladogram (Fig. 2). The Idoteidae 



GIvP'onoline group 


to Idoleinae 













LINEAGE A teduclion of 
amennal flagella 




LINEAGE B no reduction of 
antennal flagella 


2+1 1+2 



Cdolea Enchsonella 







Idolea Engidolea Glyptidolea 

1 Paridotea Synischia 

2+0 Pentias 


Colidolea Synisoma 







FiGL'RE 3. Schematic representation of possible phylogenetic pathways for pleonal fusion in the Idoteinae. 
Pleonal formulas are written above pleonal diagrams and indicate number of free and number of fused 
pleomeres. Genera assigned to given pleonal morphology are indicated below diagrams. See text for discussion. 

Stands out most strikingly from the 4 families that comprise its sister-group in the 
retention of numerous plesiomorphic attributes (e.g., free penes retained in 4 of the 5 
subfamilies; free cephalon; one pair of appendix masculina. on the second pleopods). 
Racovitza and Sevastos (1910) long ago recognized the primitive nature of the Idoteidae. 
regarding it as an '"ancient" family. The Idoteidae was the first valviferan family to be 
described, subsequent families being distinguished from it by elucidation of new char- 
acters acquired outside the Idoteidae. Thus, historically a diagnosis of the Idoteidae 
has been developed largely upon absence of characters (a phenomenon common among 
older taxa). The discovery of new distinguishing attributes (apomorphies) for the Hol- 
ognathidae and Idoteidae is clearly needed and will provide important data for testing 
the hypotheses contained in the cladogram. 

The Xenarcturidae and its sister-group (Arcturidae-Amesopodidac-Pseudido- 
theidae)are distinguished by the following synapomorphies: ( 12) first pleopods of males 
bearing "accessory appendix masculina" (in addition to the true appendix masculina 
of the second pleopods); (13) cephalon fused medially to pereonite I (second thorac- 
omere); (14) peduncle of first pleopods greatly elongated. The Xenarcturidae is a mono- 
typic taxon distinguished by the following autapomorphies: (15) pereonites I-IV with 
lateral margins expanded into large plates covering bases of pcreopods; (16) second 
antennae with flagella reduced to single articles; and (17) flagella of second antennae 
directed towards mouth, rather than away from buccal field. 

The Arcturidae and its sister-group (Amesopodidae-Pseudidotheidae) are distin- 
guished by a synapomorphy in the functional grouping of the pcreopods (18). Only in 
these taxa are pcreopods I-IV similar and directed forward to form a functional group 


distinct from pereopods V-VII. In all other valviferan taxa the percopodal functional 
grouping is I-III and I V-VII. Arcturidae is distinguished by two synapomorphies: (19) 
the unique body shape (cylindrical or tubular, often geniculate), and (20) having per- 
eonite IV generally manifestly enlarged or elongated. 

The Amesopodidae and Pseudidotheidae are distinguished by the synapomorphic 
condition of having pereonites II-IV grossly enlarged (21). Amesopodidae is a mono- 
typic family {A. richardsonae Stebbing, 1905) distinguished by the autapomorphies of 
highly reduced second pereopods (22), and the complete loss of pereopods III and IV 
(23). Pseudidotheidae contains two genera distinguished by the synapomorphy of fusion 
of the first two articles of the peduncle of the second antennae (24). 

The Family Idoteidae and the Subfamily Idoteinae 

The systematic history of the family Idoteidae can be traced through the following 
works: H. Milne Edwards (1840), Dana (1853), Bate and Westwood (1868), Harger 
(1880), Miers (1881). Chilton (1890), Ohlin (1901), Richardson (1905a), Stebbing 
(1905), Collinge (1917), Barnard (1920), Nordenstam (1933), Menzies (1950fl), Shep- 
pard (1957), and Menzies and Miller ( 1972). The American idoteid fauna is well known, 
largely due to the work of Dana (1853), Harger (1880), Benedict (1897), Richardson 
(1899a. /), 1900, 1901, 1904, 1905a. b, 1909), Hatch (1947), Menzies and Waidzunas 
(1948), Menzies (1950a. b), Menzies and Bowman (1956), Menzies and Frankenberg 
(1966), Menzies and Miller (1972), and Brusca and Wallerstein (1977, 1979a./)). All 
species of Idoteidae are marine, although two species of the subfamily Mesidoteinae 
also extend their distributions into fresh water. Saduna {=Mesidotea) entoniofi has 
been found in several deep Scandinavian lakes, and Austridotea lacustris^ occurs from 
the littoral zone to fresh water rivers and lagoons in New Zealand. 

The subfamily Idoteinae contains 22 valid genera (Table 1). The great majority 
are shallow-water and, for the most part, intertidal species. Few species are restricted 
to depths greater than 30 m. For the past 150 years (since the work of Brandt 1833 
and H. Milne Edwards 1840) studies on this group have consistently found that the 
use of a few clearly defined characters provided a basis for a classification that has been 
both stable and reliable. Thus most idoteine genera are clearly defined, unambiguous, 
and easily distinguished from one another. As will soon be seen, however, not all 
idoteine genera can be defined by unique apomorphies. The few genera that are not 
clearly differentiated from one another comprise 2 small groups of largely monotypic. 
Southern Hemisphere genera that are in need of reexamination. Principal characters 
used to distinguish the idoteine genera are external and easily recognized, as follows. 

Uropods. — Tht uropods of Idoteinae are either biramous or uniramous. The prim- 
itive biramous condition, while being clearly distinct from the uniramous condition 
and hence useful in pattern analysis, is not understood ontogenetically (see Caiman 
1909, Racovitza and Sevastos 1910, Tait 1917, Nordenstam 1933, and Menzies and 
Miller 1972). Loss of one uropodal ramus has occurred at least twice among the val- 
viferan families, in the Idoteidae (subfamily Idoteinae) and again in a single species of 
Arcturidae {Microarcturus digitalis Nordenstam 1933). Whether or not these separate 
losses were by the same "mechanism" is not known. 

Pleon. — Isopod taxa are characterized by varying degrees of fusion of the pleomeres 
and telson. Although trends towards fusion of pleomeres are evident throughout the 
Isopoda and occur in every suborder, no one has yet attempted to analyze these mor- 
phoclines in a systematic fashion (Fig. 3). In the Valvifera, there are always four or 
fewer free pleomeres, plus the pleotelson. The term "pleotelson" refers to that region 
of the pleon consisting of the telson and its fused pleomeres. For many years, the pattern 
of discrete character states manifested by fusion of pleomeres in the idoteine genera 
has been taken to represent a morphocline that is a sequence of phenotypes presumed 
to reflect the probable evolutionary pathway of descent. The polarity or direction of 
this morphocline is clearly shown by out-group analysis to be towards levels of in- 
creasing pleomere fusion. Fusion of the pleomeres is often (but by no means always) 
indicated by the presence of partly free lateral margins, distinguishable by the presence 


Table 1. Summar\- of "traditional characters" used to difFcrentiate the genera of Idoteinae (from Menzies 
and Miller 1972; with corrections). See text for additional characters. Edotca includes the synonym Ilpclys. 
and Zcnobiana includes the synonym Cleantis. Erichsonella includes the synonyms Erichsonia Dana and 
Ronalca Men/ies and Bowman. Pircs (pers. comm.) has a manuscript in preparation in which she intends 
to remove the monot> pic genus Ronalca from synonymy with Erichsonella. based on the alleged presence 
of a single pair of lateral incisions in R. pseudoculata (Boone). I have not examined R. pseiidoculata myself 
Such a change would require a minor revision in the cladogram (Fig. 17b). by adding Ronalca as a fork 
at the lip of the line leading to Eusynmicrus. making these two genera sister-taxa. 



am of Antenna 2 









Unira- Bira- 







mous mous 

Barnaididotca Menzies & Miller 


1 +0 



Crabyzos Bate 


+ 3 



Engidolea Barnard 


1 + 2 



Glyptidotca Stebbing 


+ 3 



Moplisa Morcira 


0+ 1 



Pentias Richardson 


+ 3 



Pandotca Stebbing 


1 + 2 



Synidotea Harger 


0+ 1 



Synischia Hale 


+ 3 



Idolea Fabricius 


2+ 1 



Colidotea Richardson 


0+ 1 



Synisoma Collinge 



2 + "" 



Zenobianopsis Hale 


4+ 1 • 



Euidotea Collinge 


+ 3 



Cleantiella Richardson 


1 +2 



Erichsonella Richardson 





Eusynmicrus Richardson 


0+ 1 



Parasymmerus Brusca & Wallenstein 


0+ 1 



Lyidolca Hale 


+ 3 



Zcnobiana Risso (as Zcnobia) 



3+ 1 



Cleantioides Kensley & Kaufman 


3+ 1 



Edotca Guerin-Meneville 


0+ 1 



of lateral incisions (generally referred to as "suture lines'"). Presence of free lateral 
margins on the fused pleomeres is taken to represent incomplete fusion (i.e., medial 
fusion only) of these somites, and hence a less derived state than absence of the free 
lateral margins (i.e., complete fusion). 

Maxi//ipech. — The palp of the idoteid maxilliped is composed of 5 or fewer articles. 
Out-group analysis indicates the plesiomorphic state (occurring in all families except 
Idoteidae) is 5 free articles; reduction in the number of articles thus represents a derived 
condition. It is not known with certainty whether reduction in the number of articles 
is due to fusion or to actual loss, although Brusca and Wallerstein ( 1 979c/) have suggested 
that both processes may exist among various genera (e.g., loss in Idotca, fusion in 

Anwnnac — In valviferans, the flagellum of the second antenna is either (1) nuii- 
tiarticulate (the primitive condition); (2) reduced to a single clavate article with 1 to 4 
minute "vestigial" apical articles; (3) reduced to a single clavate article only; or (4) 
reduced to just the minute "vestigial" articles. Brusca and Wallerstein i\979a) point 
out that these two kinds of reduction (clavate vs. vestigial articles) arc probably the 
result of two different processes, the former being a case of fusion of the tlagellar articles, 
the latter being an actual loss or reduction in the number of articles, creating a "vestigial" 
flagellum. In one genus (Zenobiaua) both the clavate condition and or the vestigal 
condition may both occur, suggesting that the two morphologies are somehow linked, 
perhaps both dcvelopmentally and phylogenetically. The early reduction was apparently 
a situation in which partial fusion of the flagellar articles produced the clavate mor- 


phology while retaining a few vestigial, unfused, apical articles (as seen in some 7.en- 
ohiana species). This condition could have progressed in either of two directions — loss 
of the vestigial articles to leave just the remaining clavate process (as in Cleantiella, 
Erichsonella, Euysninierus, Parasyinnieriis, Cleantioides, and Lyidotea), or loss of the 
clavate process to leave just the remaining vestigial articles {Edotea and some species 
of Zenobiana). The relationships between the clavate and vestigial conditions may not 
be resolvable in a phylogenetic sense, and may represent differing avenues of a flexible 
developmental program. However, unlike the pleonal and maxillipedal palp charac- 
teristics (above) which show varying degrees of reduction, the antennal flagella are 
either reduced (states 2-4 above) or not reduced. 

Co.xal plates. Although the coxal plates have been used extensively in valviferan 
taxonomy, use of these structures has not been consistent. Previous workers have treated 
these structures in a variety of ways and one worker's description is not always com- 
parable to another's. These problems have been discussed at length by Nordenstam 
(1933), Sheppard (1957), and Brusca and Wallerstein (1979a). For these reasons, the 
coxal plates are not considered in the following analysis. 

While the genera of the Idoteidae appear to be reasonably well-defined, the 5 
nominate subfamilies are not. The subfamily Idoteinae stands apart from the other 4 
in numerous features, and appears to represent a monophyletic group. The other 
subfamilies (Glyptonotinae, Chaetilinae, Parachiridoteinae, and Mesidoteinae) cannot 
be easily separated from one another, nor be distinguished unambiguously in a clado- 
gram. For this reason, these 4 subfamilies collectively are herein considered an out- 
group of the Idoteinae. They may be thought of as representing an unresolved poly- 
chotomy on the cladogram in Fig. 4. In the following discussion these 4 subfamilies 
are treated as one and referred to as the "glyptonotine-group.""* A second out-group 
used to construct a cladogram of Idoteinae genera is the Holognathidae (see Fig. 2). 
Numbers in the following discussion correspond to that on the cladogram in Fig. 4. 

The glyptonotine-group is distinguished by the following synapomorphies: (1) ce- 
phalon strongly produced laterally, moving eyes to dorsal position; (2) body broadened 
and dorsoventrally depressed; (3) pereopods I-III subchelate or prehensile. It retains 
the symplesiomorphy of separate penes. The Idoteinae is distinguished by the following 
synapomorphies: (4) reduction of the pleon to the 3+1 condition; (5) fusion of the 
penes into a single structure (Fig. 4). 

Paleontological data, limited as they are, corroborate the out-group comparison 
for the Idoteinae. The oldest known idoteid fossils are referred to the monotypic genus 
Proidotea {P. haugi Racovitza and Sevastos, 1910), from mid- to late Oligocene deposits 
of eastern Europe. This genus clearly falls within the subfamily Mesidoteinae (the 
glyptonotine-group). The only other fossil data for the Valvifera are Pleistocene to 
Recent specimens of Saduria {=AIesidotea)— probably the holarctic 5". sabini (Kroyer). 
In both of these genera, the pleon is composed of 4 somites, plus the pleotelson. The 
maxillipedal palp of Saduria is 5-articulate. The uropods of Mesidoteinae are biramous, 
as in the subfamilies Glyptonotinae, Chaetilinae, and Parachiridoteinae. 

The genus Zenobianopsis Hale, 1946 is not indicated on the cladogram (Fig. 4). 
The status of this deep water taxon is uncertain. Only two species have been reported 
and they differ markedly in morphology. Species of Zenobianopsis have a pleon of 4 
free somites, plus indications of a fifth (although in Z. caeca Hale, 1946, these somites 
appear somewhat reduced). Other attributes indicate that Zenobianopsis is a very 
primitive member of Idoteidae (Table 1), presumably with its origin at or about the 
time of separation of the Idoteinae from the glyptonotine-group. The early isolation of 
these species is further suggested by the fact that both are known only from deep 
subantarctic waters of the Indian Ocean. Zenobianopsis is indicated by a dashed line 
in the evolutionary tree in Fig. 1 6; further research may place the two species in separate 

Within each of the two principal idoteine lineages depicted in Fig. 4, the trend 
towards fusion of pleomeres is expressed in a "directed'' fashion. That is, the greater 
the degree of pleomere fusion in a taxon, the farther up the cladogram it appears. Fig. 















FiGi'RE 4. Cladogram of Idoteinae genera. Only apomorphies are indicated. Numbering of characters cor- 
responds to discussion in text. For characters uniting Holognathidae to Idoteidae see Fig. 2 and text. 

3 provides a schematic representation of known pleonal morphologies beginning with 
the 4-segmented pleon found in the glyptonoline-group. The pleonal formulas are 
written in two digits, separated by a plus sign. The first digit is the number of complete 
pleomeres present in the pleon (not counting the pleotelson); the second digit is the 
number of lateral incisions present, representing remnants of incompletely fused pleo- 

There are 14 possible combinations or pleonal formulas that species of Idoteinae 
might possess: 3+1. 3+0, 2+2, 2+1. 2+0, 1+3, 1+2, 1 + 1, 1+0. 0+4. 0+3. 0+2. 0+1. 
0+0. However, there are over 100 possible different steps in which fusion may progress 
to give rise to these 14 combinations. Each of these 100+ pleonal morphologies is 
derived in a unique manner and hence each constitutes an '"attribute" (scnsii Platnick. 
1979:542). Not every pathway is represented in Fig. 3: only enough steps are shown 
in order to reach the existing morphologies of the known idoteine genera in a parsi- 
monious fashion. This large ''uncertainty" problem, as well as the seeming parallelism 
of pleonal fusion, can be resolved by careful examination of the schematic pattern 
analysis in Fig. 3. All known genera of Idoteinae have pleonal formulas that must have 
arisen from one of two main lines: one line beginning with a 3+ 1 configuration (indicated 
on Figs. 3 and 4 as "lineage A"), and the other beginning with a 2+1 configuration 
(indicated on Figs. 3 and 4 as "lineage B"). Assignment of genera to one or the other 
of these lineages can be based on the antennal features described above. That is. the 
13 genera with multiarticulate second antennal flagella arc hypothesized to represent 
a lineage or series distinct from the 8 genera that have lost the multiarticulate conditions. 
Any other assignment of these genera requires accepting convergent evolution of these 
antennal morphologies; parsimony is maintained by presuming these antennal mor- 
phologies to have arisen only once. This split clearly places all genera on one or the 
other of these two main lines of descent involving pleomcrc fusion, and further suggests 
that evolution of identical pleonal formulas in these two lines of descent was through 
different steps, and hence not true convergence at all. 

Not all 14 possible patterns are represented by extant species. In fact, only 9 are 
known at present (4+1.3+1.2+1. 2+2. 1 +2. 1 +0. 0+3. 0+ 1 . 0+0). Other morphologies 

presumably existed in the past (in extinct intermediate taxa) or exist at present but 
await discovery. Fig. 3 predicts the nature (overall morphology) of these yet to be 
discovered genera, and hence sets the stage for testing the hypotheses contained therein. 

Fig. 3 is not a cladogram OR an evolutionary tree. It is merely a graphic arrange- 
ment of pleonal morphologies in a sequence of most primitive at the top, witfi increas- 
ingly derived morphologies (increased fusion) expressed following the arrows down 
and across. As can be seen in Figs. 3 and 4, reduction in pleomere number corroborates 
the reduction in the antennal flagellum (i.e., there are no conflicts). 

Lineage B (Fig. 4) is distinguished by two synapomorphies: (6) pleonal fusion has 
advanced to the 2+ 1 morphology, and (7) the uropods have lost the primitive biramous 
morphology, deriving a uniramous condition. Lineage A is defined by the synapo- 
morphy of (8) antennal flagellum reduction to condition 2 described above (i.e.. fusion 
of most flagellar articles into a clavate process bearing on its apex a few remaining 
"vestigaf" articles). 

Within lineage B, the genus Idotea is distinguished by the apomorphy of (9) re- 
duction in maxillipedal palp article number; the sister-group to Idotea has reduction 
of pleonal morphology beyond the 2+1/2+0 condition (10). Baruardidotea, Moplisa, 
and Synidotea have maxillipedal palp reduction to 3 articles (1 1), as well as continued 
fusion of pleomeres to produce a 1+0 pleonal morphology (12). Baruardidotea is 
distinguished from Moplisa and Synidotea by retention of the 1+0 pleonal formula, 
while the latter two genera have achieved the 0+ 1 condition (13). Moplisa and Synidotea 
can be distinguished from one another by the loss of the molar process in the former 

The Paridotea-Engidotea line is distinguished from its sister-group by retention 
of the symplesiomorphic pleonal condition, 1+2 (vs. the synapomorphic condition, 
0+3) (15). These two genera cannot be clearly distinguished from one another on 
morphological criteria as they are currently understood. The Colidotea-Synisoma group 
is distinguished from its sister-group by two synapomorphies: (16) lacinia mobilis of 
mandible greatly enlarged (as large or larger than incisor process), and (17) reduction 
to a 4-jointed maxillipedal palp. Colidotea is distinguished by (18) a 0+1 pleonal 
morphology. Synisonia is distinguished by two synapomorphies: (19) pleonal mor- 
phology 0+0, and (20) an elongate pleon ('/3 or more total body length). Euidotea is 
distinguished from its sister-group by (21) possession of a 4-jointed maxillipedal palp 
(vs. 5-jointed in the Glyptidotea-Pentias-Crabyzos-Synischia line). The latter 4 genera 
cannot be distinguished by morphological criteria as they are currently understood and 
are hereafter referred to as the Glyptidotea-group. 

Within lineage A, Zenohiana can be distinguished only by the symplesiomorphous 
retention of biramous uropods: whereas its sister-group has achieved the synapomor- 
phic condition of uniramous uropods (22). Some species of Zenohiana have lost the 
minute apical articles on the tip of the second antennae, while others retain these 
terminal articles. I am in agreement with previous authors that these differences do 
not warrant the splitting of Zenohiana into several genera (e.g.. Bate and Westwood 
1861-1868, Issel 1913, Collinge 1917, Barnard 1925). I have not taken the presence 
or absence of the "vestigial" articles into consideration in construction of the cladogram. 

Cleantioides, as defined here, contains only two species, C. occidentalis (Richard- 
son) and C. planicaitda (Benedict). This genus is distinguished by 2 synapomorphies: 
(23) reduction of the fourth pair of pereopods to nonambulatory appendages, and (24) 
reduction of the maxillipedal palp to 4 articles in one species (C occidentalis). It retains 
the symplesiomorphic 3+ 1 pleonal morphology, while its sister-group has achieved the 
2+2 or less stage (25). 

Cleantiella is distinguished by the apomorphy of pleonal reduction to the 1+2 
condition (26). Lyidotea and its sister-group bear 2 synapomorphies: (27) the pleon 
has achieved the 0+3 morphology, and (28) the maxillipedal palp has been reduced to 
the 3- or 4-articulate condition. Lyidotea bears an apomorphic condition in which the 
last perconal somite has become fused with pleomere 1 (29); its sister-group is distin- 
guished by the 0+2 or less pleonal morphology (30). 


Erichsonella is distinguished by the apomorphy of (31) complete pleonal fusion, 
resulting in the 0+0 condition, although some species retain a faint trans\erse furrow 
presumably representing the fused juncture of pleomere 1 to the remainder of the pleon. 
Erichsonelld's sister-group {Eiisymmerus-Parasytnmerus-Edotea) has achieved the 0+ 1 
morphology (32). Edotea-Parasyninierus are defined by the synapomorphy of maxil- 
lipedal palp reduction, to the 3-articulate condition (33). Edotca is distinguished by its 
acute subtriangular pleon (34). Eusyninicriis is distinguished by the unique 3-pointed 
spine that arises between the lacinia and incisor process on the left mandible (35). 

This cladogram (Fig. 4) reduces the number of character convergences to two. 
Reduction in the number of articles on the maxillipedal palp appears to take place time 
and again, throughout the various idoteine lines, and no sensible cladogram design can 
eliminate it. In Fig. 4 it occurs four times in lineage A and three times in lineage B. 
For this reason, it was given lower priority than all other characters used in the analysis. 
Maxillipedal palp reduction is common throughout the order Isopoda and represents 
a kind of convergence or parallelism known as canalized evolutionary potential. This 
character also expresses both "inside" and "outside" parallelism (sensu Brundin 1976. 
1981). The second convergence (or parallelism) is loss of the ancestral biramous uropod 
condition, which takes place in (and characterizes) lineage B, and then again in the 
Cleantioides through Eusymmerus line of lineage A (synapomorphy 22). 

While the cladogram in Fig. 4 is highly parsimonious, what is clearly needed is 
the elucidation of additional synapomorphies to further test the contained hypotheses. 
Six terminal taxa (or groups) cannot be distinguished by unique apomorphies at this 
time. As Sheppard (1957) and Brusca and Wallerstein (1979a) pointed out, a critical 
examination of the nature of the pereopodal coxae in the various idoteid genera will 
surely prove enlightening and undoubtedly provide us with a new suite of character 
states with which to test evolutionary relationships among the genera. The exact nature 
of the reduction in flagellar articles on the second antennae needs to be investigated 
(particularly regarding Zenobiana, Cleantioides, and Edotea), and this too might shed 
new light on the cladogram. This reduction, as well as reduction in maxillipedal palp 
articles and free pleomeres, tends to follow Brown's (1965) "Rule of Evolutionary 
Reduction." Finally, two unresolved polychotomies exist within the Idoteinae that can 
be resolved only by synonymizing genera or by the elucidation of new characters to 
differentiate these taxa. It is my belief that a careful study of the coxal plates, lacinia 
mobili. second antennae, and pleopods 3-5 could resolve all of these problematic areas 
among the idoteine genera. 


Idoteids, like other isopods, brood their developing young in a marsupium. from 
which they are hatched as "mancas," which are subjuveniles that resemble miniature 
adults except for lacking the seventh pair of pereopods. There is no evidence, ecological 
or morphological, that the manca stage is planktonic. and in those species that have 
been reared in the laboratory, hatchlings are always benthic crawlers like their parents. 
The only published records of idoteids in plankton are for the 2 widespread species, 
Cleantioides planicaiida and Idotea metallica, and the 2 species belonging to the ques- 
tionable genus Zenohianopsis. Both mancas and adults of most species, however, are 
capable of swimming in short bursts over small distances (e.g., between algal fronds). 
When idoteids are dislodged from the substratum by waves or surge they sink or swim 
quickly to the bottom (Jansson and Matlhiesen 1971, Salemaa 1979, Sywula 1964. Lee 
and Gilchrist 1972. Wallerstein and Brusca 1982). Idoteids are clearly a component of 
the benthic community in marine shallow-water habitats, where their niche is that of 
a cryptic herbivore and occasional scavenger. This suggests that idoteids, particularly 
intertidal species, are probably not good dispersers. 

Because the following discussion deals with the Idoteinae at the generic level, it 
must remain fairly general. Even at this level, however, these generalizations describe 
patterns of geographic distribution that can be correlated with the cladogram (Fig. 4), 

1 1: 

Figure 5. Distribution of valviferan families (excluding Idoteidae): Amesopodidae [O], Arcturidae [•], 
Holognathidae [D], Pseudidotheidae [O], Xenarcturidae [A]. 

and which can most parsimoniously be ascribed either to vicariance or non-vicariance 
events. Geological dates of events described in this section have been used to transform 
the cladogram into the evolutionary tree depicted in Fig. 16. It is not my purpose to 
present a detailed analysis of each genus here; that must await a species-by-species 
study of each genus (e.g., see Brusca 1983 for the genus Colidotea). The latter will 
clearly be an enormous undertaking, but one for which the following analysis could 
provide a starting point. 

The Valvifera as a whole show strong correlation to southern temperate latitudes, 
3 of the 6 families being restricted to that geographic region, 1 known only from the 
shores of India, and the other 2 being widespread (Fig. 5). The idoteine genera show 
a similar geographic trend (Figs. 7-14). Numerous studies have recently appeared that 
examine the relationships of distributional patterns of terrestrial flora and fauna to the 
geological history of the Southern Hemisphere (e.g., Keast 1973, Raven and Axelrod 
1972, Cracraft 1974, 1980). However, similar studies on marine groups have been few. 
For these reasons, Gondwanan shores are discussed in some detail. A brief review of 
the breakup of Gondwana follows, based on Kennett (1977), Smith and Briden (1977), 
Norton and Sclater (1979), Knox (1979), Durham (1979), Grant-Mackie (1979) and 
Hallam (1981). 

Throughout most of the Permian, the Triassic, and most of the Jurassic periods, 
Gondwana was unified as a single land mass. The proto-southern continents were all 
situated at higher latitudes than they are today. Although Permian glaciation probably 
existed in the highest southern latitudes, that cold era came to a fairly abrupt halt by 
the Jurassic. By mid-Mesozoic times climates in the Southern Hemisphere had changed 
markedly, as a long-lasting global warming trend became established. Paleontological 
evidence suggests that by the middle or late Jurassic the northern shores of Gondwana 
had already begun acquiring a warm-water Tethyian biota, thus restricting temperate 
coastal biota to the southernmost latitudes. A common temperate marine fauna prob- 
ably inhabited the contiguous coastline of Gondwana south of 55°-65°S latitude. This 
shallow coastal sea washed the shores of southernmost South America, southeast Africa, 
eastern Antarctica and eastern Australia. The continuous coastal topography, coupled 
with non-glacial and generally equable climates of the Jurassic, facilitated *'warm-water 
cosmopolitanism" along the shores of northern Gondwana. and "cold-water cosmo- 
politanism" along the shores of southern Gondwana. These two regions were probably 
physically isolated from one another until at least the early Tertiary, when separation 
of Australia from Antarctica instituted a direct high latitude southern connection be- 
tween the Indian and Pacific oceans. The distribution of modern coastal temperate 


faunas in the Southern Hemisphere can thus be hypothesized to be. at least in part, 
the product of the fragmentation of an early to mid-Mesozoic temperate Gondwanan 

That part of Gondwana composed of Antarctica, Australia, New Zealand and South 
America rotated southward during the Mesozoic, although as early as the Late Jurassic 
these land masses were beginning to separate as Antarctica-Australia-India began un- 
coupling from Africa and South America. By the Early Cretaceous (about 120 MY A) 
sea floor spreading had started to form the south Atlantic Ocean. At about the same 
time, India began to decouple from Antarctica-Australia-Africa. Marine conditions 
developed between India and Antarctica- Australia by 105 MY A. Africa was clearly 
separated from Gondwana about 90 MYA. New Zealand split from Antarctica-Aus- 
tralia 70-80 MYA. By the mid-Cretaceous the south Atlantic Ocean was open along 
its entire length as Africa and South America pulled away from one another. The free 
ocean connection (the "Vema Gap") between the north and south Atlantic was estab- 
lished by the Late Cretaceous as the transverse Rio Grande-Walvis Ridge sank below 
the 1 km depth (approximately 78-80 MYA). Australia was last to decouple from 
Antarctica, about 50-60 MYA. During the Paleocene these land masses were probably 
separated by a shallow narrow seaway; however, the South Tasman Rise acted as an 
effective barrier to the development of a circum-Antarctic current until about 30-50 
MYA, subsequent to which the southern circum-polar current began to develop. The 
modern deep-flowing Antarctic Circumpolar Current could not have been established 
until the opening of the Drake Passage, 22-28 MYA, when deep-sea conditions de- 
veloped between southern South America and Antarctica (plate boundaries and timing 
of geotectonic events in this region are still very controversial). 

During most of the course of events described above, Antarctica continued a slow 
drift southwards, reaching a position close to its present location by Late Cretaceous: 
it has remained nearly stationary throughout the Cenozoic. Thus, deep marine con- 
ditions began to develop in the Southern Ocean by the Late Paleocene, although a 
corridor of scattered highlands and shallow seas persisted between Australia and Ant- 
arctica until about the Late Eocene or Early Oligocene. 

South America has moved progressively westward since the Late Cretaceous. India 
collided with Asia about this same time. Coincidental with this southern fragmentation, 
the highest post-Carboniferous sea levels ever to occur (Campanion Era, about 75 
MYA) submerged roughly 'A of the present continental area below shallow epiconti- 
nental seas. Deep water flows from the Pacific into the Atlantic across Central America 
commenced about 55 MYA, only to be halted in the Pliocene when the Panama Isthmus 

The shallow-water marine fauna would not have responded to the breakup of 
Gondwana in the same manner as the terrestrial fauna, because the initial separation 
of the land masses created intervening shallow seaways that fostered the spread of 
marine biota before separation of the biotas occurred (for a comparison with the 
fragmentation of the terrestrial fauna of Gondwana see Cracraft 1 974 and Rosen 1 978). 
As South America and Africa moved northward, and Antarctica moved southward, 
relative to one another, the effect on temperate coastal marine life was most likely to 
have created two disjunct faunas. One of these faunas, the western coastal fauna, would 
have inhabited the shores of southern South America, southern Africa, and north- 
western Antarctica. The eastern fauna would have been restricted to Australia, eastern 
Antarctica and southernmost India. New Zealand, at this time, was situated in warm- 
water latitudes. By the mid-Cretaceous, the Southern Ocean had become extensive 
enough to break the western fauna into two separate temperate regions: southern South 
America (and probably the region of the Antarctic Peninsula) and southern Africa. The 
eastern Gondwana. mid-Cretaceous, temperate coastal region remained unchanged 
even though New Zealand had split from the warm-water shores of Australia. By the 
mid- to Late Cretaceous the shallow sea between India and Antarctica had deepened 
to isolate these regions entirely from one another. India having drifted into the tropical 
waters of the Indian Ocean. The coastal environment of southern Australia remained 



"& ;f 

* A® 







So. America 

s. Australia, 

e. Antarctica 
l^ India 




So. America, Africa, 
nw. Antarctica 

s. Australia, New Zealand, 
e. Antarctica, India 


Figure 6. Proposed geographic cladogram of temperate, shallow-water, Gondwanan coastal areas. Note 
the significant differences between this cladogram of shallow continental shores (marine geographic areas) 
and Rosen's (1978) cladogram of Gondwanan continents (terrestrial geographic areas). Seelexl for discussion. 

tied to northern Antarctica until the Cretaceous-Tertiary boundary, or perhaps even 
to the Early Oligocene. These relationships are pictured in Fig. 6. 

Because paleoclimates and oceanographic conditions were not the same during the 
Mesozoic and Cenozoic as they are today, latitudinal distributions of shallow-water 
marine life would have been under quite different thermal regimes. This ecological 
component must be taken into account when considering paleobiogeographical recon- 
structions. A brief review of Mesozoic-Cenozoic climates follows; for more detailed 
information consult Fleming (1975, 1979), Margolis et al. (1978), Grant-Mackie ( 1 979). 
Durham (1979), Frakes (1979), Zinsmeister (1982), and references therein. 

Overall, middle and late Mesozoic climates were extremely equable, with ocean 
temperatures exceeding those of the present day by 7°-15°C. Polar ice caps did not 


Figure 7. Composite distribution map of all idoteine genera, except the 3 cosmopolitan taxa (Idoiea. 
Synidotea, Zenobiana). Dashed lines indicate limits of warm-water (tropical/subtropical) regions; open cir- 
cles = genera of lineage A; closed boxes = genera of lineage B. 

exist and both terrestrial and marine biotas exhibited little evidence of latitudinal 
zonation. Cosmopolitanism was common. Cool sea water existed in the southern oceans 
only south of about 50°S latitude. Early Cretaceous sea bottom temperatures were 
approximately 10°-15°C warmer than today; Late Cretaceous sea bottom temperatures 
were approximately 7°C warmer than today. The Eocene marked the beginning of a 
global cooling trend that ultimately led to the Cenozoic glaciations and cooling of deep 
ocean bottom waters to their present thermal regimes (Shackleton and Kennett 1975). 
As late as the Eocene (38-55 MYA), all but the southernmost Antarctic seas were 
warm-temperate in nature. The steep thermal drop over the Eocene/Oligocene boundary 
was probably related to the establishment of the Antarctic Circumpolar Current and 
subsequent South Polar glaciation. Although land masses occupied both poles by the 
Early Paleocene, large-scale Antarctic glaciations probably did not begin until the Mio- 
cene. The Antarctic Convergence began moving northward in the Oligocene. Cooling 
trends continued throughout the Cenozoic, apparently marked by several periods of 
severe chilling. 

Fossil reefs, requiring relatively warm surface waters, grew to paleolatitudes of 
about 40° throughout the Paleogene, as in the Cretaceous. In the northeast Pacific, 
tropical environments (i.e., surface temperatures >20°C) extended to approximately 
45°N in the Eocene, and have contracted equatorward steadily since that time. Abun- 
dant paleoceanographic and stratigraphic data exist to indicate that New Zealand shores 
were bathed primarily by warm subtropical waters throughout the Cretaceous and 
Paleocene (Fleming 1962, 1975, Durham 1979, Knox 1979), although a distinct cooling 
trend began in the middle to Late Miocene, eventually resulting in present-day tem- 
perate coastal conditions. At best, only southernmost New Zealand might have expe- 
rienced temperate waters from the Eocene to the Miocene. By early Pleistocene sub- 
antarctic waters had reached about 40°S latitude on New Zealand shores. The middle 
Oligocene marked the end of "'high latitude subtropical communities," and by the Late 
Miocene modern water bodies and their associated biological provinces had begun to 
be established. The Australian Subantarctic water mass had formed by about 1 5 MYA. 
For the past 10-15 million years the Southern Ocean temperature, circulation and 
zonational water masses have remained essentially the same. 

If the distributions for all species of Idoteinae are plotted on the globe, nearly every 
sea and shore is seen to be inhabited by one or more genera. Fig. 7 is a composite 
distribution map plotting the ranges of all taxa of Idoteinae EXCEPT the 3 cosmopolitan 
gQnQV2i — Idotea, Synidotea. and Zenobiana. These 3 taxa are omitted from the com- 


Figure 8. Distributions of Cleantiella [•], Lyidotea [•], and Cleantioides [O]. 

posite distribution map because they provide no particular insights into a global pattern 
analysis at this level. Distribution maps of the individual genera are given in Figs. 8- 
14. and are discussed below. It will be seen from these distribution maps and the 
following discussion that most idoteine genera are endemic to only one or a few coastal 
regions. The dashed lines on Fig. 7 indicate the limits of the tropical/subtropical regions, 
based on Ekman (1953), Briggs (1974), and Brusca and Wallerstein (1979/)). While 
some disagreement exists regarding the exact limits of the tropical zones, the latitudes 
that I have chosen clearly separate the "warm-water" faunal regions of the world's 
shores from the "cold-water" regions. The only serious arguments with these delimi- 
tations might be in whether one regards the Mediterranean Sea as warm-temperate or 
subtropical. The best comprehensive discussion of Mediterranean zoogeography is 
probably that of Ekman (1953), who in describing the complex mixtures of northern 
and southern elements, couldn't comfortably label this sea either warm-temperate or 
subtropical. The nature of south African faunal designations has also been a matter of 
some controversy, and was recently reviewed by Brown and Jarman (1978). 

Several general patterns are revealed when Fig. 7 is examined. In the New World, 
endemic idoteine genera are distributed fairly equally in both warm and cold waters. 
In the Old World, however, there is a marked absence of records from the tropics. 
Only 3 genera on this map represent Old World tropical taxa: Cleantiella and Clean- 
tioides both occur on the east Asian coast, while Lyidotea is known from a single species 
in northeastern Australia (see Fig. 8). A second pattern seen is that Old World endemic 
genera tend to form 3 distinct clusters, as follows: (1) a Southern Hemisphere cold- 
water group, (2) a European cool- to cold-water group, and (3) a northeast Asian cool- 
to warm-water group. Only 2 of the genera depicted in Fig. 7 occur in both the Old 
and New Worlds: the southern temperate Paridotea and the northern tropical Clean- 
tioides. Because most species and genera of Idoteidae are restricted to temperate or 
polar seas, the family has long been considered a "cold-water centered taxon" (see 
Brusca and Wallerstein, 1919b and references therein). It is of particular interest that 
the Old World and New World tropical regions are inhabited by entirely different 
genera. The New World tropical genera are Cleantioides. Erichsonella, Parasyninierus, 
and Eusymmerus. The Old World tropical genera are Cleantiella, Lyidotea, Idotea, 
Synidotea, and Zenohiana. As the latter 3 are largely cosmopolitan taxa, the obvious 
question is, "Why have species in these genera been unable to invade the tropics in 
the New World?" Let us now examine the distributions of the individual idoteine 

Idotea is nearly a cosmopolitan genus (Fig. 9). It occurs in most waters of the Old 
World, including the tropics, but in the New World it is notably absent from the warm 
seas between the Tropics of Cancer and Capricorn in both the Pacific and Atlantic. 


Figure 9. Distribution of Idotea. 

Brusca and Wallerstein ( 1 919b) reviewed literature documenting the eurythermal nature 
o{ Idotea species, and suggested that biological factors (competition or predation), rather 
than temperature, could be excluding the 2 New World temperate isopod genera, Idotea 
and Synidotea, from the Western Hemisphere tropics. Wallerstein and Brusca (1982) 
subsequently provided experimental and comparative morphological evidence that 
predation by crustacivorous fishes is possibly restricting species o^ Idotea from the New 
World tropics. In the latter study they proposed a model that envisioned "faunal waves" 
of tropical predator species (primarily fishes) moving northward from the equatorial 
region during warm Pleistocene interglacials, pushing the southern latitudinal range 
end points of non-adaptable temperate species northward ahead of them, and thus 
excluding Idotea and Synidotea from the New World equatorial region (see Addicott 
1 970 and Zinsmeister 1 974 for a discussion of similar north-south faunal displacements 
in Mollusca). The nearly global distribution of Idotea, as well as its position on the 
cladogram (Fig. 4) relative to other genera in lineage B (see discussion below) suggests 
that Idotea is an old genus (pre-Cretaceous). It should be emphasized that, while the 
genus Idotea is cosmopolitan, the species in this taxon are themselves largely endemic 
to restricted coastlines. Only one species of Idotea is cosmopolitan, /. nieta/llca. Its 
cosmopolitanism has been explained by drift dispersal of the algae on which it lives 
(Naylor 1972, Poore 1981). 

Nine of the 12 remaining genera in lineage B are endemic to the Southern Hemi- 
sphere (Table 2) and are probably descendants of a pan-austral, cold-water, Gondwanan. 
pre-Cretaceous fauna. The concept of a pan-austral terrestrial biota was apparently first 
recognized by Hooker (1853, 1860) and Huxley (1868). Though largely suppressed by 
100 years of "Wallaceian dispersalism," the concept has finally experienced a rebirth 
owing largely to the work of Pantin et al. (1960) and Brundin (1966. 1970. 1972<:/. /). 
1976). For a recent summary of many subsequent papers see N.Z. DSIR (1979). Only 
recently, however, has serious documentation and discussion of generalized temperate 
pan-austral marine tracks begun (e.g., Zinsmeister 1976, 1982, Fleming 1975. Knox 
1975). The mid-Cretaceous/early Tertiary fauna that inhabited this temperate, shallow- 

Table 2. Distribution of the 8 Idoteinae Genera of Lineage B Endemic to the Southern Hemisphere. 


















Southern South America 



South Africa 






Southern AustraHa 





New Zealand 





Figure 10. Distributions of Glyptidotea and Barnardidotea [•], Engidotea [•]. Pentias [A], Zenobianopsis 
[D], and Paridolea [O]. 

water, southern Gondwanan region has been referred to as the WeddelHan Province 
by Zinsmeister (1976). 

One of the eadiest appearing genera in this posX-Idotea radiation was Synidotea, 
which like Idotea is nearly cosmopolitan but notably absent from the New World 
tropics (Fig. 1 1 ). The same comments that apply to Idotea above are probably applicable 
to Synidotea. The origins of Synidotea appear old enough that the Early Cretaceous 
circum-Arctic seaway probably served as one of several principal dispersal routes, this 
being reflected in the modern distribution of the genus, which has by far the majority 
of its species concentrated in the Northern Hemisphere. This hypothesized age of origin, 
plus the restriction of both Barnardidotea and Mop/isa (Synidotea^s sister-group) to 
the Southern Hemisphere, argues strongly for the origin of Synidotea in the southern 
seas. This contrasts with the opinion of Gurjanova (1935) and Menzies and Miller 
(1972), who suggested that the place of origin of Synidotea was the north Pacific. 
Menzies and Miller's opinion appears to have been based solely on the fact that most 
species of Synidotea presently occur in this area, which they considered to be its "center 
of origin." The present analysis, however, does corroborate Menzies and Miller's (op. 
cit.) dating of the origin of this genus. Synidotea's sister-group, Moplisa, consists of a 

FiGURt 11. Distribution of .S'lYj/^o/f'a. 


Figure 12. Distribution of Crabyzos [•], Synischia [•], Ewdotea [D], Synisoma [O], Colidotea [A], and 
Moplisa [O]. 

single species. M. sphaewmiformis (Mane-Garzon. 1946). so far known only from a 
short stretch of coastline in southern Brazil (Fig. 1 2). The cosmopolitanism ofSynidoiea 
(vs. the restricted distribution of the monotypic Moplisa). and the absence of an apo- 
morphy to distinguish Synidotea from Moplisa strongly suggests that Moplisa evolved 
either in sympatry or as a peripheral isolate from a continuing stock of Synidotea. 
These relationships are pictured in the phylogram (Fig. 16) and are consistent with the 

Although Glyptidotea, Crabyzos, Pentias, and Synischia cannot be separated by 
cladistic analysis (Fig. 3), the biogeographic data provide evidence regarding their 
origins. Glyptidotea is endemic to temperate South Africa (Fig. 10); Crabyzos and 
Synischia are endemic to temperate Australasia (Fig. 12). These patterns suggest that 
these 3 genera arose subsequent to the initial east-west split of Gondwana (i.e.. post- 
Jurassic). The absence o{ Glyptidotea from the South American component of the west 
Gondwanan track suggests that this genus arose subsequent to the separation of these 
two land masses (i.e.. mid-Cretaceous or later). The closely related genus Pentias is 
today restricted to temperate shores of northern Japan (Fig. 10). The simplest expla- 
nation for its occurrence would seem to be fortuitous jump dispersal across the warm 

Figure 13. Distribution of Ze«oWa«a. 


Figure 14. Distribution of Eusymmerus and Parasymmerus [•], Edotea [O], and Erichsonella [•]. 

waters of the equatorial region, perhaps during a period of global cooling and tropical 
compression such as the late Tertiary (although the western Pacific was probably far 
less affected by marine cooling than other regions on the globe). A second, remote 
possibility is that it (or its ancestors) reached Japan via China, which may have originally 
been part of the Gondwanan continent (see Crawford 1974). 

None of the above southern taxa occur along the warmer shores of the Indian 
Ocean. Their distributions suggest that these genera are Cretaceous to post-Cretaceous 
descendants of the widespread Jurassic-Early Cretaceous temperate Gondwanan track 
discussed earlier in this section. The ancestor(s) inhabiting this track is indicated in the 
evolutionary tree by "GonAnc" (Fig. 16). 

Because several genera cannot be clearly distinguished from one another by the 
cladogram, it is difficult to seek congruence between lineage B and the geographic 
cladogram of Fig. 6. However, a partial correlation (38%) is nonetheless evident (Fig. 
\5a, b). Table 2 provides a distributional summary of the 8 Idoteinae genera of lineage 
B that are Southern Hemisphere endemics. Fig. 15 gives reduced taxon-area and geo- 
graphic cladograms for the groups in question. The cladogram congruence with the 
Synidotea-Baniardidotea-Moplisa line corroborates the hypothesis that the latter two 
taxa arose subsequent to the splitting of South America from Africa, while Synidotea 
continued to persist unchanged. Correlation also exists for Synischia. If one accepts 
the probable dispersal of Crahyzos from Australia to New Zealand, the correlation 
between these cladograms is raised to 50%. Paridotea has retained its circumpolar 
distribution, corroborating its early (pre-Cretaceous) origin. 

The-occurrence of Euidotea on African, Australian, and New Zealand shores (Fig. 
12), and its absence from South American coasts today, argues for either (1) its origin 
prior to the break-off of Africa and its subsequent extinction in South America, or (2) 
its origin on African shores after the break-off of that continent and its subsequent 
spread to Australasia via West Wind Drift. Recall that New Zealand probably did not 
achieve its present temperate coastal thermal regime until well after the deep water 
barriers were formed that isolated it from Australia, probably not until the Eocene at 
the earliest. Thus, introduction of the cold-water genus Crahyzos could have been via 
dispersal from Australia in fairly recent times (mid- to late Tertiary). This same rea- 
soning must apply to the other temperate New Zealand genera, Paridotea and Euidotea. 
The fact that none of these three taxa are endemic to New Zealand (all 3 also occur in 
southern Australia) lends support to this probability. 

The sister-groups Colidotea and Synisoma are restricted to warm-temperate and 
subtropical waters of the New and Old World respectively (Fig. 12). This restriction, 
plus the widespread occurrence of Synisoma (Europe, Far East), suggests that these 











— 3 

ID d) 
O 2 



0) ~ 
• ^ -H 
•H U-J 


- C 

ID 10 

O M 

■H ID 

1-1 UJ 

M-l N 


ID 2 

< ID !-^ 













x; -u 






JJ 01 







3 3 




• —  




m < 


















— ' 































































Figure 1 5a. Reduced geographic cladogram of 4 extant temperate shores derived from Gondwana {see Fig. 
6). 15b. Reduced taxon-area cladogram of Southern Hemisphere endemic idoteine genera (see Fig. 3 and 
Table 2). 

taxa are vicariant descendants of a widespread, circumglobal. northern Tethyian track. 
The absence of either of these genera on eastern American shores is most easily ex- 
plained by local extinction on that coast. Closure of the Panama seaway in the Pliocene, 
or simple isolation from the parental stock as the North Atlantic opened up. could 
have provided the setting for the origin of these two taxa as we know them today. 
Judging by its extreme disjunct distribution. Synisoma has remained unchanged since 
at least the Paleocene, when the Gibralter gap closed. Thus, it appears that the ancestors 
of these two taxa were members of the Tethyian fauna, at least marginally, while the 
extant descendants have been pushed northward to the subtropical and warm -temperate 
portions of their former range. 

In summary, the genera comprising lineage B appear to have remained closely tied 
to their ancestral (Jurassic) temperate Gondwanan distribution. Historical hypotheses 
that are consistent with the cladogram suggest that vicariancc alone can account for 
the modern distributional patterns of only a few taxa. most haxing modern distributions 
clearly tied to both former vicariant and dispersal/extinction events. These probable 
extinctions, dispersal events, and biological interactions affecting ranges add to the 
uncertainty in ascribing geographic origins to the relatively old taxa comprising lin- 
eage B. 

Turning to lineage A. we see one broadly distributed genus, the primitive Zcno- 
biana. which occurs in both temperate and tropical waters of the Old World, but is 
restricted to temperate South America in the New World (Fig. 13). Zenobiana is the 
only genus of Idoteinae to retain the primitive biarticulatc uropods. .As was the case 
with Idotea, Zenobiana's cladistic relationship and widespread distribution initially 


"9 5; 

6 5; 

i a 5 i 

Figure 16. 

Proposed phylogeny of the genera of Idoteinae, based on Fig. 1 7 and other data. See text for 

suggests that Zenobiana was pre- to Early Cretaceous in origin (i.e.. pre-Gondwanan 
fragmentation). Zenobiana cannot be much older than this as it clearly shows no alliance 
to the Permo-Triassic Tethyian Sea radiation, owing to its near absence from the tropical 
Indo-West Pacific region; only one species occurs in Indo-Pacific waters. Z. nata/ensis 
(Barnard 1925). Thus. Zenobiana's limited invasion of tropical environments appar- 
ently took place after the final closure of the Tethyian Sea. in the early to mid-Tertiary. 
Its alliance to and probable origin on temperate shores is further indicated by its present 
distributional pattern. All genera that appeared subsequent to the establishment of the 
globally distributed Zenobiana (the 7 remaining genera in lineage A) are warm-water 
taxa, or at least probably initially evolved as warm-water taxa (i.e.. Edotea). The absence 
of identifiable apomorphies to distinguish Zenobiana indicates that this genus was both 
the ancestor to, and persisted after the origin of its sister-group. 

Cleantioides consists of only 2 species: C. occidentalis is endemic to the tropical 
eastern Pacific; C planicauda is a widespread tropical species known from both coasts 
of the Americas and from eastern Asia. Cleantiella and Lyidotea are western Pacific 
descendants of a Cleantioides-Mke ancestor. Cleantiella is restricted to the warm shores 
of USSR and China, and most likely arose as a post-Cretaceous northwest Pacific 
endemic (Fig. 8). Lyidotea is restricted to the warm waters of northeastern Australia. 
The absence of Cleantiella and Lyidotea, both warm-water taxa, from the equatorial 
region of the west Pacific is enigmatic and suggests 3 possibilities: (1) dispersal of the 
ancestor of Lyidotea from Asian shores to Australian shores; (2) the former existence 
of a N-S tropical Cleantiella-like track joining these two regions, with subsequent 
fragmentation into northern and southern tropical isolates with extirpation in the equa- 
torial region; or (3) an artifact of collection records. The paucity of records for species 
in any other genus of Idoteidae from this region, and the unlikelihood of alternatives 
1 and 2 above, suggest the third possibility may be the correct interpretation. According 
to Poore (in litt.) virtually no collecting has ensued in this region since the Siboga 
Expedition (ca. 1900). 


All remaining genera in lineage A are New World endemics and represent descen- 
dants of the post-Cretaceous ancestors of this line. Edotca has an anti-tropical distri- 
bution, occurring on both North and South American shores (Fig. 14). Brusca and 
Wallerstein (1979a) have discussed this unusual distribution, concluding that Edotea, 
although evolving in the New World tropics as part of the early transisthmian track 
biota (the "Tertiary Caribbean Province" of Woodring 1954, 1966; the "Panamanian 
Track" of Croi/.at et al. 1974), was promptly excluded from this warm-water region 
by competition with the better adapted tropical genera like Clcantioides, Eusymmcrus, 
Parasymmenis, and Erichsonella. Modern species of Edotea now occur only in tem- 
perate latitudes to the north and south of the New World tropics. "Better adapted" 
includes the possession of such predator avoidance adaptations as small body size, 
early reproduction, and cryptic morphologies (see Wallerstein and Brusca 1982, for 
details of these adaptations). 

Edotea. Erichsonella (Fig. 14), and Cleantioides are all components of the Tertiary 
Caribbean Province (Woodring 1966) that have retained their amphi-American dis- 
tributions. Numerous eastern Pacific-western Atlantic analog or geminate species can 
be identified within these genera. Eusymmenis and Parasymnwrus {Fig. 14) are tropical 
eastern Pacific endemics. Parasyninierus probably evolved subsequent to the Pliocene 
closure of the Panama seaway. However, if the relationships depicted on the cladogram 
are correct, Eusymmerus probably evolved prior to the closure of the seaway (in order 
to be the sister-group or ancestor of Edotea-Parasymmerus). Its present restriction to 
the eastern Pacific implies that either (1) it evolved there and never became part of the 
transisthmian fauna, or (2) it was part of the transisthmian biota but has subsequently 
become extinct in the western Atlantic. The latter seems far the more probable sequence 
of events. Woodring (1966) has pointed out that many molluscan genera presently 
restricted to the tropical eastern Pacific occurred throughout the eastern Pacific-western 
Atlantic Tertiary Caribbean Province during Miocene times. These geographic data, as 
well as the cladistic relationships, imply that Edotea was both the ancestor to Para- 
symmerus and remained essentially unchanged subsequent to the origin of Parasym- 
merus (Fig. 16). It is unfortunate that fossil material for isopods is rare, for such data 
could provide excellent corroboration or refutation of the above hypothesis.-^ 

The data summarized in the cladogram and biogeographic review clearly imply 
that there were both Old World and New World warm-water genera derived from an 
ancestral global ^^Zenobiana-Cleantioides line." The extant Old World fragments of 
this line are Cleantiella and Lyidotea\ the extant New World fragments are Erichsonella, 
Edotea, Parasymmerus, and Eusymmerus. 

The relationships in the cladogram suggest that these latter 4 New World endemic 
taxa were derived as the sister-group of Lyidotea (an Old World endemic). This ne- 
cessitates trans-Pacific (west to east) dispersal of the ancestor of these 4 New World 
genera. An alternate scenario would have the New World taxa of lineage A descended 
not from the geographically restricted Lyidotea line or its Old World ancestor, but from 
the widespread ancestral Zenobiana-Cleantioides line. This could be accomplished 
simply by reconstructing a portion of lineage A, as shown in Fig. 1 7. This new cladogram 
of lineage A is longer than the original (Fig.4) in requiring 9 pleonal transformations, 
versus 8 in the original. The new cladogram requires four transformations (but two 
convergences) in maxillipedal palp reduction, as in the original. Thus, for lineage A we 
are faced with a cladogram slightly more parsimonious (one less step) but requiring a 
major successful trans-Pacific dispersal event to establish the entire New World tropical 
lineage of idoteine genera, versus a cladogram slightly less parsimonious but requiring 
only an easily explained vicariant origin of the New World line from a pre-drift cos- 
mopolitan track. This is perhaps a case of strict methodological parsimony versus 
"biological parsimony." Given the multitude of avenues for pleonal reduction. I see 
no compelling reason to accept one cladogram over the other. Such problematic re- 
lationships can probably be resolved only by the elucidation of new generic-level at- 
tributes, and by careful examination of the morphological and biogeographic relation- 
ships of the individual species in these genera. 



The above phylogenetic and biogeographic analysis sheds light on several problems 
in understanding the evolution of the Idoteinae. First is the problem of discordant 
character trends. Although trends toward fusion of body somites and appendage articles 
(pleon, antennae, maxillipedal palp) have been noted for many decades, it was not until 
Menzies and Miller (1972) summarized and tabulated these data that the discordant 
trends in these different morphological features were recognized, suggesting probable 
high levels of homoplasy in the Idoteinae (see Table 1). Generating schematic pattern 
analyses (Figs. 4 and 17a) reveals that identical pleonal morphologies can be derived 
in a great many ways. For example, previous workers have assumed the 0+1 pleonal 
morphology of Colidotea to have been derived in the same manner as the 0+ 1 pleonal 
morphology of Eusynimeriis, Parasymnierus, and Edotea (i.e., to be homologous), 
despite the fact that the former has a multiarticulate second antennal flagellum whereas 
the latter 3 genera have uniarticulate (clavate) second antennal flagella, the latter pre- 
sumably being a fundamentally significant difference. In fact, the 0+1 pleonal mor- 
phology can be derived in any of about a dozen different ways, and in the above case 
it is clear that the derivation of Colidotea was probably quite different from that of the 
other 3 genera (see Figs. 3, 4, 16 and 17). Use of this schematic approach to pattern 
analysis in the Idoteinae also facilitates placement of the primitive genus Zenohiana 
in a single lineage apart from Cleantioides, even though both taxa have identical pleonal 
morphologies and have been confused with one another in the past. Further, the sche- 
matic analysis provides clear visual descriptions of all potential intermediate stage 
morphologies for the Idoteinae genera, lending high predictability (hence testability) 
to hypotheses contained in the above analysis. 

That isopods, and peracarids in general, were already diverse and important taxa 
by the late Paleozoic has been fairly well documented by Schram (1970, 1974). By as 
early as the Pennsylvanian, the Peracarida had radiated into most if not all its currently 
recognized orders (except perhaps the Amphipoda). As Schram (1974) stated, ". . . 
peracarid radiation was probably initiated in Devonian time, when it is generally 
thought caridoid eumalacostracans were taking origin," ". . . the Peracarida are now 
known to be among the most ancient of the eumalacostracans," and ". . . the superorder 
Peracarida was a major element in the late Paleozoic radiation of eumalacostracans 
and contributed the principal caridoid types of that time." The earliest isopods known 
from the fossil record are phreatoicids from the middle Pennsylvanian; spelaeogripha- 
cians, tanaids, and mysidaceans first appear in the lower Carboniferous. 

The origins of the subfamilies of Idoteidae hypothesized in this study are consistent 
with Schram's data, and also with dates offered by Schultz (1979), who stated that most 
isopod families were well-defined before the Triassic. 

The present analysis reveals the existence of two principal lines of descent within 
the Idoteinae (Fig. 16), lines that separated from one another early on (Jurassic or 
perhaps even Triassic). One of these lines remained closely tied to its origins in the 
Old World Southern Hemisphere temperate biota, while the other invaded New World 
environments and undertook a radiation in the New World tropics (producing genera 
such as Cleantioides, Erichsonel/a, Eusymmerus, and Parasyiuinerus). Brusca and Wal- 
lerstein (1979/?) hypothesized that idoteids might only recently have invaded the New 
World tropics, perhaps in close association with a similar invasion of these regions by 
the brown algae (Phaeophyta), which probably serve as the principal habitat and food 
resource for most idoteine species. That the late Triassic-Early Jurassic periods might 
have had a widespread southern temperate Idoteinae distribution of little or no ende- 
mism is hardly surprising. As Hallam (1981) points out, Jurassic marine faunas as a 
whole were decidedly more cosmopolitan than those of the present day. These early 
origins in temperate waters explain the long-held belief that the Idoteidae and Idoteinae 
are "cold-water centered taxa." Plate movements became increasingly influential in 
promoting endemism within the Idoteinae from Late Jurassic through Late Cretaceous, 
as Pangaea fragmented. The high degree of endemism among Idoteinae genera contrasts 


sharply with claims by Taylor and Forester (1979) that cold-water psychrospheric 
isopod biofacies (generic assemblages) tend to be similar regardless of latitude or depth. 

This study has shed some light on the place of origin of both the Idoteinae and 
the Idoteidae, as well as the Valvifera itself. Distributions of the Idoteidae subfamilies 
(excluding the Idoteinae) are given in Fig. 18; distributions of the five other valviferan 
families are given in Fig. 5. As can be seen, two of the five Idoteidae subfamilies are 
distinctly Gondwanan in distribution (Glyptonotinae and Chaetilinae). one is predom- 
intly Laurasian (Mesidoteinae), one is ubiquitous (Idoteinae). and one is endemic to 
the shores of Morocco (Parachiridoteinae). Of the six valviferan families, two are 
cosmopolitan (Idoteidae and Arcturidae), and the remaining four are all Gondwanan, 
including the most primitive family, the Holognathidae. No valviferan family is dis- 
tinctly Laurasian in distribution. These data taken together strongly suggest that the 
Valvifera originated in the temperate Southern Hemisphere at least by early or mid- 
Pangaean times (Permian/Triassic), if not earlier. 

Hurley and Jansen (1977) reviewed the zoogeography of the isopod family Sphae- 
romatidae on Southern Hemisphere coastlines. Their data on generic and species dis- 
tributions reflect patterns similar to those seen here in the Idoteinae. Hurley and Jansen 
found generic affinity between all southern continents, as well as generic endemism on 
all shores (endemism levels reported are 12 genera [48%] in Australia; 7 genera [41%] 
in South Africa; 3 genera [20%] in New Zealand; and 1 genus [1 1%] in Chile). Hurley 
and Jansen invoked strict dispersalist mechanisms to account for these distributions, 
however, and stated, "The possibility of invoking continental drift as an agency is 
hardly necessary in view of the fact that littoral species are involved.'" What this 
statement is meant to imply is not clear. Presumably the authors are suggesting that 
littoral organisms are expected to disperse across ocean barriers with great regularity, 
thus negating the roles of continental drift and vicariance in establishing endemism. 
This view seems inconsistent with their data, however, which record high levels of 
endemism at both generic and species levels on southern continental shores (species 
endemism on Australian shores was reported as 91%; South African shores, 80%; New 
Zealand, 89%; Chile, 62%). In any event. Hurley and Jansen (1977) did not present 
any phylogenetic analyses of the taxa in question, making it impossible to evaluate 
alternative biogeographic scenarios for the Southern Hemisphere Sphaeromatidae. 

The present study reveals several situations wherein ancestral taxa apparently 
persisted while new (sister) taxa evolved as peripheral isolates (or perhaps in sympatry) 
in restricted geographic regions. For example, Synidotea almost certainly persisted 
unchanged during the events that produced Moplisa and Barnardidotea. Similar situ- 
ations exist for: Paridotca and Engidotea: Zenobiana and its sister-group; and Edotea 
and Parasynifnerus (see Fig. 16). 

This study suggests that both dispersal and vicariant forces probably played im- 
portant roles in creating modern-day distributional patterns of idoteine genera. Dis- 
persal via the Antarctic Circumpolar Current appears to have played only a minor role, 
however, as endemism on southern shores is high. The unique (highly endemic) nature 
of these southern continental shores was established long ago. Even New Zealand, 
which spans 13 degrees of latitude and sits just 1760 km off Australia, is noted for its 
high endemicity of coastal species. Examples include 24% endemism for polychaetes 
(Augener 1924), 50% for crabs (Chilton and Bennet 1929), 64% for echinoderms (Mor- 
tensen 1925), and 89% for sphaeromatid isopods (Hurley and Jansen 1977). Because 
the present analysis was at the generic level rather than the species level, and because 
several Southern Hemisphere genera cannot be clearh distinguished in a cladogram. 
resolution of all geographic patterns has not been achieved. What is clearly needed are 
similar analyses for each of the idoteine genera (e.g., see Brusca 1 983). It is of particular 
interest to note that the major lines of descent within the Idoteinae appeared prior to 
the mid- to Late Cretaceous global warming trend, no doubt further facilitating the 
temperate-based distribution of this taxon. The present study adds further evidence to 
a large body of data demonstrating the concept of the Crustacea being a taxon fraught 
with convergences. As Schram (1978) deftly pointed out, "The central dominating 




Cleantiella Lyidotea 

3+1 2+2 1+3 0+4 0+3 

loss of 
biramous uropods 




retention of 
biramous uropods 























FkiIkk 17a. Alternative schematic representation for lineage A, based on biogeographic analysis. 17b. 
Alternative cladogram for lineage A, based on 17a. Synapomorphies are as follow: (1) uropods uniramous, 
(2) pereopods IV reduced, (3) maxillipedal palp reduced in some species to 4 articles, (4) pleon 2 + 2. (5) 
maxillipcdal palp reduced to 4 articles. (6) pleon 0+1. (7) pleon 1+2. (8) pleon 0+0. (9) maxillipedal palp 
reduced to 3 articles. (10) pleon with unique, acute, subtriangular shape, (11) left mandible with 3-pointed 
spine arising between lacinia and incisor, ( 1 2) pereonite VII fused to pleonite 1,(13) maxillipedal palp reduced 
to 3-4 articles, (14) pleon 0+3. See text for discussion. 


Ficii'RE 18. Distribution of the subfamilies of Idoteidae (except Idoteinae): Mesidoteinae [•]. Chaetilinae 
[a], Glyptonotinae [O], Parachiridoteinae [O]. 

theme of arthropod evolution is the muhiphcity of convergent development. No phy- 
letic scheme, monophyletic or polyphyletic, can escape this." 

Wiley (1981) divided biogeography into three subdisciplines: descriptive, histor- 
ical, and ecological biogeography. He felt that the goals and interests of the ecological 
biogeographer lie more with ecology than with systematics, whereas the opposite is 
true of the historical biogeographer. Finally, he suggested that systematics has little to 
contribute directly to the field of ecological biogeography. I disagree with Wiley, and 
feel I have shown here and through a series of studies that there is a logical sequence 
of basic taxonomy/descriptive biogeography/ecological-historical biogeography. all these 
studies existing within the realm of systematics (i.e., Brusca and Wallerstein 1977. 
1979fl, b, Wallerstein and Brusca 1982, Brusca 1983). Hessler and Wilson (in press) 
provide further evidence of these relationships by their implication of both ecological 
and historical factors in reviewing the probable causes for differences in distribution 
of major crustacean taxa. Pregill and Olson (1981) came to similar conclusions with 
regards to the Caribbean land vertebrates, as did Stock (1981) regarding the Caribbean 
crustacean stygobionts. While the methods of ecological and historical biogeography 
may differ, the two endeavors are clearly complementary; to attempt one at the expense 
of the other is to invite error. 

Only a few genera of Idoteinae can be confidently tied to vicariance events asso- 
ciated with the breakup of Pangaea. Only one New World genus {Parasymmenis) can 
be clearly tied to such major geological events. Vicariant relationships seem to be more 
easily distinguished at the species level {see Brusca 1983). It would seem that, at least 
in idoteid isopods, too much time has passed and too many unknowable events tran- 
spired (new taxa have evolved and gone extinct, extant taxa have undergone local 
extinctions in selected portions of their range, dispersal events, etc.) to confidently 
extract clear vicariant patterns at the level of supraspecific taxa. Furthermore, those 
idoteinc taxa that do appear to be products of vicariance phenomena can only be tested 
by comparison to cladograms and distributional data for other intertidal groups. Cra- 
craft (1982) has recently detailed such a procedure. Briefly, allopatric vicariant specia- 
tion (type la of Bush 1975) implies the appearance of a barrier. In the case of littoral 
isopods, these barriers would be new stretches of ocean or new land barriers across 
shallow seaways. Such a barrier would be expected to influence the vicariance patterns 
of numerous intertidal taxa. and one would thus predict that concordant pairs of sister- 
taxa would exist on either side of the barrier. In contrast, speciation resulting from a 
dispersal event to a new area (a founder individual or population; type lb speciation 
of Bush 1975) is generally taken to be a random event. Hence, one would predict not 


to find concordant vicariance patterns from one clade to another. Absence of such 
concordance with other intertidal taxa thus would suggest that speciation was initiated 
not by a vicariance event, but by a dispersal event. Thus, the need for cladograms of 
other marine invertebrate groups becomes evident, and without such studies one cannot 
critically assess the hypothesized processes responsible for the patterns present today 
in the idoteine taxa. Finally, the low levels of congruence between the cladogram of 
taxa-area and geographic cladogram can also be explained by the simple hypothesis 
that Gondwanan distributions of ancestral idoteid taxa were NOT widespread, but 
fragmented and local. This possibility, of course, denies a popular premise of generalized 
tract theory, that ancestral ranges can be assumed to be the sum of the ranges of the 


' There have been several recent attempts to cast doubt on the monophyletic nature of the Peracarida 
(Watling 1981, Dahl and Hessler 1982, Hessler in press). This is not the place to present detailed analyses 
and criticisms of each of these studies. However, it should be pointed out that in Watling's "cladistic" analysis 
only certain selected characters were used, numerous character transformations were illogical and unexplained, 
character states attributed to various taxa were incorrect, and a clear method of character polarity assessment 
was not provided. Although the other recent studies (op. cit.) employed a variety of approaches, often in the 
guise of cladograms, none attempted to analyze character state polarities in a strict logical order or with any 
clearly expressed methodology. Schram (1981) was concerned with the recognition of basic structural plans 
within the Eumalacostraca, and his classification is based on a strictly random array of character associations. 
The phenogram he chose for conversion into a classification is the one he felt "most comfortable with." 
Hessler's (in press) "cladogram" of the Peracarida (his fig. 5) is entirely unjustified by the data he presents. 
Among other problems, no attempt was made to achieve parsimony and the "cladogram" actually requires 
more convergences (at least 24) than its contained character transformations (23). In the case of Watling 
(1981) and Hessler (in press) it appears as though evolutionary scenarios were conceived first, these then 
being transformed into dendrograms (incorrectly called cladograms) upon which the appropriate "apomor- 
phies" were overlain. Hessler's dendrogram of the Peracarida is essentially Siewing's (1963 and earlier 
publications) concept of peracarid relationships. This procedure, of course, ignores parsimony considerations 
and is the exact reverse of what a phylogenetic (cladistic) analysis is meant to accomplish. 

The fact is. there exist many unique synapomorphies that unite the peracarid orders: (1) maxilliped with 
basis produced into an anteriorly directed, bladelike endite; (2) lacinia mobilis present in adults; (3) oostegites 
formed on inner pereopodal coxal margin; (4) young brooded in a brood chamber or "marsupium" (the 
location of the brood chamber varies from a simple oostegial pouch to invaginations of the sterna, modified 
oviducts, or even to the inner carapace region in the thermosbaenaceans); (5) direct development, with no 
true postnaupliar larval stages; (6) release of the young as "manca"; (7) whiplike immobile sperm, devoid 
of fibrils and mitochondria (this character needs further documentation); (8) a large suite of embryological 
attributes {see below); and (9) a large suite of attributes associated with the functional morphology of the 
pereopods. These functional and morphological skeletomuscular adaptations are associated with a system 
unique to the Peracarida, in which the body-coxa articulation has lost the caridoid "gimbal" joint and become 
either immobilized or capable of only limited abduction/adduction, and the coxa-basis articulation become 
monocondylic but capable of performing a complete suite of motions. This peracaridan system for ambulation 
IS present in the incipient condition in mysidaceans. Hessler (1982) views these particular peracaridan leg 
synapomorphies as adaptations necessitated by the development of the marsupium and its attendant oos- 
tegites. Although thermosbaenaceans have lost the oostegites, they still retain the infolded monocondylic 
articulation of the pereopodal coxa and basis {see Hessler 1982. for details). 

The loss of the oostegites in the thermosbaenaceans is probably a response necessitated by unique 
morphological (fusion of pereopodal articles) and functional (locomotory) adaptations of the pereopodal 
endites in this group. Loss of oostegites and concomitant relocation of the brood chamber is not unique to 
the thermosbaenaceans among the Peracarida (it occurs in several hyperiid amphipods and isopod higher 
taxa). This matter has been competently dealt with by Fryer (1964) and need not be repeated here (also see 
Hessler 1982 and Slewing 1958). 

The presence of lacinia-like movable spines in the larvae of a few species of euphausids and shrimps 
suggests two possibilities: (1) the lacinia of adult peracarids is a paedomorphic attribute {sensu Gould 1977) 
retained from a nonperacaridcan ancestor with lacinia in larval stages only, or (2) these are superficially 
similar convergent features. The absence of postnaupliar larval stages in the Peracarida, and their direct 
development to a juvenile hatching stage, argue for their origin not from a eucarid ancestor but from a line 
separate from the eucarids. Hence, the second hypothesis is the more parsimonious. The structural simplicity 
of the "lacinia" of eucarid larvae, versus the complexity of the peracarid lacinia supports this contention. 
Should one choose to consider the movable mandibular spine of certain adult bathynellaceans to be true 
lacinia (see Siewing 1963, Schminke 1972, and Dahl and Hessler 1982), one is confronted again with two 
possibilities: (1) the adult lacinia is a feature representing a potential synapomorphy for a bathynellacean- 
peracarid line, or (2) this character is convergent in these two groups. I would accept either interpretation 
as a working hypothesis. The latter seems the more likely considering the presence of movable spines in 
certain eucarid larvae and the apparent plasticity of the spines of the mandibular row. The point is, however, 
that acceptance of either hypothesis would not affect the monophyletic status of the Peracarida! Frankly, 


with regards to the monophyly of the Peracarida. all the fuss over the lacinia seems "much ado about 

The Peracarida retain an "underlying unity in development" (Anderson 1973) distinct in numerous ways 
from the Eucarida and Syncarida. This unity has been well documented and succinctly summarized by 
Anderson (1973). Unique attributes of pcracaridan development appear in virtually all stages of embryo- 
genesis, and include among other things: the early segregation of primordial germ cells as a definite pre- 
sumptive area; the teloblastic development of the postnaupliar segments: the vitellophage modifications in 
the early development of the midgut, the unique embryogeny of the digestive glands: the distinct cmbryogeny 
of the ectoderm; and the formation of a second pair of "dorsal organs" (the ectodermal dorsolateral organs). 

The "mancoid" stage appears to be little more than the product of alterations in embryogeny and timing 
in the release of the young. Its absence in mysids and amphipods may be tied to a more rapid embryological 
development (or to a delayed postembryonic development) in these two taxa {see Steele and Steele 1975), 
which may also be linked to the presence of ventrally curved embryos and completion of cleavage in the 
early stages in these groups (i.e., rapid early holoblastic cleavage). Furthermore, although amphipods leave 
the marsupium with all 7 pairs of pereopods "in place." there is a great deal of variation in the structure 
and development of this appendage, particularly in the hyperiids. Laval (1980) has even recognized hyperiid 
"larvae," with distinct hatching stage morphologies. Much of the distinction between Laval's "larvae" and 
the adults involves the nature of the pereopods, and some hyperiids certainly appear to have "virtual 
mancoids." The seventh pereopods seem especially plastic in hyperiids and amphipods in general. The 
rudimentary nature of the seventh pereopods in "juvenile" amphipods of certain species has been known at 
least since the work of Bate (1861) on Vihilia. One could also posit the origin of the mancoid stage subsequent 
to the origin of the mysids and amphipods during peracaridan cladogenesis. This would remove this attribute 
from the list of synapomorphies defining the Peracarida, but it would certainly not destroy the monophyletic 
nature of that ta.xon; rather, it would simply make the "mancoid stage" a synapomorphy defining a subset 
within the Peracarida. 

An embryological attribute that might suggest alliance among the amphipods, mysids and eucarids is 
the retention (from the larval stages) of the functional antennal glands in these three taxa. Since all Crustacea 
have antennal glands during their embryogeny, the retention of these glands into adulthood in these taxa 
hardly seems surprising and is most likely either a convergence or simply a symplesiomorphy retained from 
a common ancestor (i.e., a plesimorphy not lost until the appearance of the mysids and amphipods had been 
achieved during peracaridan cladogenesis). 

I believe that a carefully (and correctly) accomplished cladistic analysis of the Peracarida will reveal the 
amphipods to be the nearest relative if not the sister-group of the isopods. No such analysis has been published 
to date, although one is in preparation by F. Schram. Watling (1981), Hessler (in press) and others have 
chosen to ignore or deemphasize the fundamental synapomorphies unique to these two taxa (e.g., sessile 
compound eyes; pereonites with coxal plates: pereopods entirely uniramous; carapace entirely lost; mandible 
of the transverse biting type), and rely instead on differences and retained plesiomorphies in their analyses. 
As indicated in Fig. 2, I do not believe that there are ANY shared derived characters unique to the tanaids- 

- Note that were the amphipods taken to be the sister-group to the isopods, attributes 1, 2 and 4 would 
become synapomorphies uniting these two taxa. 

' The genus Aiistndotea is in need of further study; it may have to be removed to the Idoteinae. 

■* Poore (in litt.) has suggested that the 4 non-idoteine subfamilies are synonymous and should be 
combined. At the time of this writing I am in general agreement with Poore, but do not address the matter 

'• The single Old World species of Edotea (E. dilatata Thomson, 1884) has been shown to be the female 
of Crabyzos elongatus (Miers 1876) (see Hurley 1961:292). 


A number of people took time to read and criticize numerous early drafts of this 
paper. The additional data and discussions they provided me were invaluable to the 
successful completion of this study, and I am indebted to them for their sincerity and 
effort: J. L. Barnard, T. Bowman, N. Bruce, G. Brusca, P. Delaney, J. Garth. R. Hessler, 
J. Haig, E. Iverson, B. Kensley. P. McLaughlin. M. Miyamoto. A. M. S. Pires. J. Savage, 
and especially F. Schram. G, Pregill. and G. Poore. The philosophical perspecti\es of 
these reviewers more often than not differed markedly from one another, and I hope 
that I have succeeded in attending to each of their criticisms in a fair and appropriate 
manner. However, the views expressed in this paper are. of course, my own respon- 
sibility. I also want to thank the former students of my graduate seminars in systcmatics 
and biogeography for their often highly emotional discussions on the subjects addressed 
in this paper, particularly E. Iverson, M. Miyamoto, B. White, and B. Wallerstein. 
Many people assisted me in obtaining specimens on loan: I especially want to thank 
the Invertebrate Zoology staff of the British Museum (Natural History). G. Wilson. W. 
Cooke (NOSC-Hawaii Lab). T. Bowman, and G. Schull/. Marie Hoff-Steinauer exe- 
cuted the art work (Exhibits Department, San Diego Natural History Museum). This 
study was funded by a grant from the National Science Foundation (DEB 80-17835). 


Literature Cited 

Addicott. W. O. 1970. Tertiarv' paleoclimatic 
trends in the San Joaquin Basin, California. U. 
S. Geol. Survey, Professional Pap. 644-D. 19 

Anderson, D. T. 1973. Embryology and Phylog- 
eny in the Annelids and Arthropods. Pergamon 
Press, New York. 495 pp. 

Augener, H. 1924. Polychaete von den Auckland- 
und Campbell-Inseln. Vid. Meddel. Dansk Na- 
turhist. Foren. 75:1-1 16. 

Barnard, K. H. 1920. Contributions to the crus- 
tacean fauna of South Africa. No. 6. Further 
additions to the list of marine Isopoda. Ann. 
So. African Mus. 17(1 1):3 19-438. 

Barnard, K. H. 1925. Contributions to the crus- 
tacean fauna of South Africa. No. 9. Further 
additions to the list of Isopoda. Ann. So. Af- 
rican Mus. 20(5):381-410. 

Bate, C. S. 1861. On the morphology of some 
Amphipoda of the division Hyperina. Ann. 
Mag. Naturl. Hist. (Ser. 3) 8:1-16. 

, and J. O. Westwood. 1861-1868. A His- 
tory of the British-Sessile-Eyed Crustacea. 
London. John Van Voorst, [in 23 parts; 1 Oct 
1861-31 Dec 1868] 536 pp. 

Benedict, J. E. 1897. A revision of the genus Syn- 
idotea. Proc. Acad. Natural Sci. Philadelphia 

Brandt, J. F. 1833. Conspectus monographie 
Crustaceorum Oniscordorum Latreilli. Bull. 
Soc. Imp. Nat. Moscou 6:171-193. 

Briggs, J. C. 1974. Marine Zoogeography. Mc- 
Graw-Hill, New York. 475 pp. 

Brown, A. C. and N. Jarman. 1978. Coastal ma- 
rine habitats, in J. Werger (ed.). Biogeography 
and Ecology of Southern Africa, Vol. II. Junk, 
Lochem, Holland. 1434 pp. 

Brown, Jr.. W. L. 1965. Numerical taxonomy, 
convergence and evolutionary reduction. Syst. 
Zool. 14:101-109. 

Brundin, L. 1966. Transantarctic relationships and 
their significance, as evidenced by the chiron- 
omid midges, with a monograph of the 
subfamily Podonominae, Aphrotaenidae and 
the austral Heptagiae. Kungliga Svenska 
Vetenskapsakademiens Handlingar 4(11):1- 

. 1970. Antarctic land faunas and their his- 
tory. Pp. 42-53 in M. M. Holdgate, (ed.). Ant- 
arctic Ecology, Vol. 1. Academic Press, New 

. 1 972a. Circum-Antarctic distribution pat- 
terns and continental drift. 1 7th Internat. Zool. 
Cong., Theme 1:1-11. 

. \912b. Phylogenetics and biogeography. 

Syst. Zool. 21:69-79. 

. 1976. A ncocomian chrionomid and 

Podonominae-Aphroteiinae (Diptera) in the 
light of phylogenetics and biogeography. Zoo- 
logica Scripta 5(3-4): 139-160. 
Brusca, R. C. 1 983. Two new idoteid isopods from 
Baja California and the Gulf of California 
(Mexico) and an analysis of the evolutionary 
history of the genus Colidotea (Crustacea: Is- 

opoda: Idoteidae). Trans. San Diego Naturl. 
Hist. Mus. 20(4):69-79. 

— , and B. R. Wallerstein. 1977. 'The marine 
isopod crustaceans of the Gulf of California. I. 
Family Idoteidae. Amer. Mus. Novitates, No. 

— , and B. R. Wallerstein. 1979a. The marine 
isopod crustaceans of the Gulf of California. 
II. Idoteidae. New genus, new species, new rec- 
ords and comments on the morphology, tax- 
onomy and evolution within the family. Proc. 
Biol. Soc. Wash. 92(2):253-271. 

and B. R. Wallerstein. 1979/). Zoogeo- 

graphic patterns of idoteid isopods in the 
northeast Pacific, with a review of shallow-water 
zoogeography for the region. Bull. Biol. Soc. 
Wash. 3:67-105. 

Bush, G. L. 1975. Modes of animal speciation. 
Ann. Rev. Ecol. Syst. 6:339-364. 

Caiman, W. T. 1 909. A Treatise on Zoology. Part 
VII. Appendiculata, Crustacea. R. Lankester 
(ed.). London. Adam and Charles Black. 346 
pp. [reprinted by A. Asher and Co., Amster- 
dam, 1964]. 

Chilton, D. 1890. Revision of the New Zealand 
Idoteidae. Trans. New Zealand Inst. 22:189- 

Cracraft, J. 1 974. Phylogeny and evolution of the 
ratite birds. The Ibis 1 16(4):494-541. 

. 1980. Biogeographic patterns of terrestrial 

vertebrates in the southwest Pacific. Paleo- 
geogr., Palaeoclim., Paleoecol. 31:353-369. 
-. 1982. Geographic differentiation, cladis- 

tics, and vicariance biogeography: reconstruct- 
ing the tempo and mode of evolution. Amer. 
Zool. 22(2):4 11-424. 

Chilton, C, and E. W. Bennett. 1929. Contribu- 
tion for a revision of the Crustacea Brachyura 
of New Zealand. Trans., Proc. New Zealand 
Inst. (Wellington) 59:731-778. 

CoUinge, W. E. 1917. A revision of the British 
Idoteidae, a family of marine isopods. Trans. 
Roy. Soc. Edinburgh 60:721-760. 

Crawford, A. R. 1974. A greater Gondwanaland. 
Science 184:1179-1181. 

Croizat, L., G. Nelson, and D. E. Rosen. 1974. 
Centers of origin and related concepts. Syst. 
Zool. 23(2):265-287. 

Dahl, E., and R. R. Hessler. 1982. The crustacean 
lacinia mobilis: a reconsideration of its origin, 
function and phylogenetic implications. Zoo. 
J. Linn. Soc. 84:133-146. 

Dana, J. 1853. On the geographical distribution 
of Crustacea. Pp. 696-805 in United States' 
Exploring Expedition during the years 1838- 
1842, under the command of Charles Wilkes, 
U.S.N., Vol. 14, No. 2. Crustacea. Pp. 696- 

de Jong, R. 1980. Some tools for evolutionary 
and phylogenetic studies. A. Zool. Syst. Evo- 
lut.-Forsch. 18:1-23. 

Durham, J. W. 1979. The fossil record, plate tec- 
tonics, and development of characteristic 


members of austral marme faunas. Pp. 165- 
186 in N. Z. DSIR [cited below]. 

Ekman, S. 1953. Zoogeographv of the Sea. Lon- 
don. Sidgwick and Jackson. 417 pp. 

Eldredge, N.. and J. Cracraft. 1979. Introduction 
to the symposium. Pp. 1-5 //; J. Cracraft, and 
N. Eldredge (cds.). Phylogenetic .Analysis and 
Paleontology. Columbia Univ. Press, New 
York. 233 pp. 

, and . 1980. Phylogenetic 

Patterns and the Evolutionary Process. Meth- 
od and Theory in Comparative Biology. Co- 
lumbia Univ. Press, New York. 349 pp. 

Endler, J. A. 1982. Problems in distinguishing 
historical from ecological factors in biogeog- 
raphy. Amer. Zool. 22:441-452. 

Felsenstein, J. 1973. Ma.ximum-likelihood esti- 
mation of evolutionary trees from continuous 
characters. Amer. J. Hum. Genet. 25:47 1-492. 

Fleming, C. A. 1 962. New Zealand biogeography: 
a paleontologist's approach. Tuatora 10(2):53- 

. 1975. The geological history of New Zea- 
land and its biota. Pp. 1-86 //; G. Kuschel (ed.). 
Biogeography and Ecology in New Zealand. 
Monogr. Biol. 27. W. Junk. The Hague. 

1979. The Geological History of New 

Zealand and Its Life. Auckland Univ. Press. 
141 pp. 

Frakes. L. A. 1979. Climates Throughout Geo- 
logic Time. Elsevier Scientific Publishing Co., 
N.Y. 310 pp. 

Fryer, G. 1964. Studies on the functional mor- 
phology and feeding mechanisms of Monodella 
argentani Stella (Crustacea: Thermosbaena- 
cea). Trans. Roy. Soc. Edinburgh 66:49-90. 

Gould. S.J. 1977. Ontogeny and Phylogeny. The 
Belknap Press. Harvard Univ. Press, Cam- 
bridge. 501 pp. 

Grant-Mackie,J. A. 1979. Cretaceous-Recent plate 
tectonic history and paleoceanography devel- 
opment of the Southern Hemisphere. Pp. 27- 
42 in N. Z. DSIR [cited below]. 

Gurjanova, E. F. 1935. Zur zoogeographie de 
Crustacea Malaconstraca des Arktischen ge- 
bietes. Zoogeographica 2:555-571. 

Hallam, A. 1981. Relative importance of plate 
movement, eustasy and climate in controlling 
major biogeographical changes since the early 
Mesozoic. Pp. 303-330 in G. Nelson, and D. 
E. Rosen (eds.). Vicariance Biogeography — A 
Critique. Columbia Univ. Press. New York. 
593 pp. 

Harger, O. 1880-1881. Report on the marine Is- 
opoda of New England and adjacent waters. 
Rpt. U.S. Comm. Fish & Fisheries, 1878, Pt. 
6:297-462. Washington, D.C. 

Harper, C. W., Jr. 1979. A Batesian probability 
view of phylogenetic systcmatics. Sysl. Zool. 

Hatch. M. H. 1947. The Chelifera and Isopoda 
of Washington and adjacent regions. Univ. 
Wash. Publ. Biology 10(5): 1 55-247. 

Hennig, W. 1966. Phylogenetic Systemalics. 
Translated by D. Dwight Davis and Ranier 
Zangerl. Univ. Illinois Press. Urbana. 263 pp. 

Hessler, R. R. 1982. The structural morphology 

of walking mechanisms in cumalacostracan 
crustaceans. Phil. Trans. Royal Soc. London 
(Ser. B) 296: 245-298. 

— . in press. A defense of the caridoid facies; 
wherein the early evolution of the Eumalacos- 
traca is discussed. Pp. 145-164 in F. R. Schram, 
(ed.). Crustacean Phylogeny. Balkema Pubis., 

— , and G. D. Wilson, in press. The origin 
and biogeography of malacostracan crusta- 
ceans in the deep sea. Spec. Publ. Syst. Assoc. 
(London). [Proc. Internat. Symp. on Biogeogr. 
"Time and Space in the Emergence of the Bio- 
sphere" London, 1981] 

-, and D. Thistle. 1979. The deep- 

sea iospods: a biogeographic and phylogenetic 
overview. Sarsia 64:67-75. 

Hooker, J. D. 1853. The botany of the Antarctic 
voyage of H. M. Ships Erebus and Tenor in 
the years 1 838-1 843. Pt. II. Flora Novae-Zeal- 
andiae. Vol. 1. Lovell Reeve. London. 312 pp. 

. 1860. On the origin and distribution of 

species: introductory essay to the flora of Tas- 
mania. Amer. J. Sci". & Arts (2)29:1-25, 305- 

Hull, D. L. 1979. The limits of cladism. Syst. 
Zool. 28(4):4 16-440. 

Hurley, D.E., and K. P. Jansen. 1977. The marine 
fauna of New Zealand: Family Sphaeromati- 
dae (Crustacea: Isopoda: Flabellifera). Mem. 
New Zealand Oceanogr. Inst. 63:1-95. 

Huxley, T. H. 1868. On the classification and 
distribution of the Alectoromorphae and Het- 
eromorphae. Proc. Zool. Soc. London, 1868: 

Issell. R. 1913. RicherchedeetologiesullTsopodo 
tubicola Zenobiana prismatica (Risso). Arch. 
Zool. Exper. Gen. 51:450-479. 

Jansson. A.-M., and A.-S. Matthiesen. 1971. On 
the ecology of young Idotea in the Baltic. Pp. 
71-88 //; D. J. Crist (ed.). Fourth European 
Mar. Biol. Symposium, Cambridge Univ. Press. 
599 pp. 

Keast, A. 1973. Contemporary biota and the sep- 
aration sequence of the southern continents. 
Pp. 309-343 in D. H. Tariing, and S. K. Run- 
corn (eds.). Implications of Continental Drift 
to the Earth Sciences. Vol. 1. Academic Press, 
New York. 622 pp. 

Kennett. J. P. 1977. Cenozoic evolution of Ant- 
arctic glaciation, the circum-Antarctic Ocean, 
and their impact on global paleoceanography. 
J. Geophysical Res. 82(27):3843-3860. 

Kitts. D. B. 1981. (Review of . . .) N. Eldredge 
and J. Cracraft. Phylogenetic Patterns and the 
Evolutionary Process: Method and Theory in 
Comparative Biology. 1980. Columbia Univ. 
Press. Science 210:1239-1240. 
Knox, G. A. 1975. The marine benthic ecology 
and biogeography. Pp. 353-403 m G. Kuschel 
(ed.). Biogeography and Ecology in New Zea- 
land. Monogr. Biol. 27. W. Junk, The Hague. 

. 1979. Distribution patterns of Southern 

Hemisphere marine biotas: some comments 
on their origins and evolution. Pp. 43-82 m 
N.Z. DSIR [cited below]. 
Laval. P. 1980. Hyperiid amphipods as crusta- 


cean parasitoids associated with gelatinous 
zooplankton. Oceanogr. Mar. Biol. Ann. Rev. 

Lee, W. L., and B. M. Gilchrist. 1972. Pigmen- 
tation, color change and the ecology of the ma- 
rine isopod Idotea resecata (Stimpson). J. Exp. 
Mar. Biol. Ecol. 10:1-27. 

Margolis, S. V., P. M. Kroopnick, and D. E. Good- 
ney. 1978. Cenozoic and late Mesozoic pa- 
leoceanographic and paleoglacial history re- 
corded in circum-Antarctic deep-sea sediments. 
Mar. Geo!. 25:131-147. 

McDowall, R. M. 1978. Generalized tracks and 
dispersal in biogeography. Systematic Zool. 

Menzies, R. J. 1950a. The taxonomy, ecology and 
distribution of northern California isopods of 
the genus Idothea with the description of a new 
species. Wasmann J. Biol. 8(2): 155-1 95. 

. \95Qb. A remarkable new species of ma- 
rine isopod, Erichsonella crenulata n. sp., from 
Newport Bay, California. Bull. So. Calif. Acad. 
Sci. 49:29-35. 

, and T. E. Bowman. 1956. Emended de- 
scription and assignment to the new genus 
Ronalea of the idotheid isopod Erichsonella 
pseudoculata Boone. Proc. U.S. Nat. Mus. 

, and D. Frankenberg. 1966. Handbook on 

the Common Marine Isopoda of Georgia. Univ. 
Georgia Press, Athens. 93 pp. 

, and M. A. Miller. 1972. Systematics and 

zoogeography of the genus Synidotea (Crus- 
tacea: Isopoda) with an account of California 
species. Smithsonian Contrb. Zool. No. 102: 

, and R. J. Waidzunas. 1948. Postem- 

bryonic growth changes in the idopod Penti- 
dotea resecata (Stimpson), with remarks on their 
taxonomic significance. Bio. Bull. 95: 107-1 1 3. 

Miers, E. J. 1881. Revision of the Idoteidae, a 
family of sessile-eyed Crustacea. J. Linn. Soc. 
London 16:1-88. 

Milne Edwards, H. 1840. Histoire Naturelle des 
Crustaces, comprenant 'anatomic, la physiol- 
ogic et la classification de ces animaux. Li- 
brairie Encyclopedique de Roret, Paris 3:120- 

Morse, J. C. and D. F. White, Jr. 1979. A tech- 
nique for analysis of historical biogeography 
and other characters in comparative biology. 
Syst. Zool. 28(3):356-365. 

Mortensen, T. 1925. Echinoderms of New Zea- 
land and the Auckland-Campbell Islands, III- 
V. Vid. Medd. Dansk Naturh. Foren 79:261- 

Naylor, E. 1972. British Marine Isopods. Aca- 
demic Press, London. 86 pp. [Synopses of the 
British Fauna, n. ser.. No. 3, The Linnean Soc. 

Nelson, G. and N. Platnick. 1981. Systematics 
and Biogeography. Columbia LJniv. Press, New 
York. 567 pp. 

Nordenstam, A. 1933. Marine Isopoda of the 
families Scrolidae, Idolheidae, Pseudidothei- 
dae, Arcturidae, Parascllidae, and Stentriidae 
mainly from the South Atlantic. Further Re- 

sults of the Swedish Antarctic Exped., 1901 — 
1903 3(l):l-284. 

Norton, I. O., and J. G. Sclater. 1979. A model 
for the evolution of the Indian Ocean and the 
breakup of Gondwanaland. J. Geophysical Res. 

N.Z. DSIR. 1979. Proceedmgs of the Interna- 
tional Symposium on Marine Biogeography and 
Evolution in the Southern Hemisphere. Auck- 
land, New Zealand. 17-20 July 1978. Vol. 1 & 
2. New Zealand Dept. Sci. Indust. Res. (N.Z. 
DSIR), Information Ser. No. 137. 

Ohlin, A. 1901. Isopoda from Tierra del Fuego 
and Patagonia. I. Valvifera. Svenska Esped. till 
Magellandslanderna 2(1 1):26 1-306. 

Pantin, C. F. A., et al. 1960. A discussion on the 
biology of the southern cold-temperate zone. 
Proc. Royal Soc. London 152B:429-682. 

Platnick, N. I. 1979. Philosophy and the trans- 
formation of cladistics. Syst. Zool. 28:537-546. 

Poore, G. C. B. 1981. Marine Isopoda of the Snares 
Islands, New Zealand— 1. Gnathiidea, Valvi- 
fera, Anthuridea, and Flabellifera. New Zea- 
land J. Zool. 8:331-348. 

Pregill, G. K., and S. L. Olson. 1981. Zoogeog- 
raphy of West Indian vertebrates in relation to 
Pleistocene climatic cycles. Ann. Rev. Ecol. 
Syst. 12:75-98. 

Racovitza, E. F., and R. Sevastos. 1910. Proidotea 
haugin g., n. sp. Isopode Oligocene de Rou- 
manie et les Mesidoteini Nouvelle Sousfamille 
des Idotheidae. Arch. Zool. Exper. Gen. 46(5): 

Raven, P. H., and D. I. Axelrod. 1972. Plate tec- 
tonics and Australasian paleogiogeography. 
Science 176:1379. 

Richardson, H. R. 1899a. Key to the isopods of 
the Pacific coast of North America, with de- 
scriptions of twenty-two new species. Proc. U.S. 
Nat. Mus. 21:815-869. 

. 1899/). Key to the isopods of the Pacific 

coast of North America, with descriptions of 
twenty-two new species. Ann. Mag. Nat. Hist. 
4(7):157-187, 260-277, 321-338. 

. 1900. Synopsis of North American in- 
vertebrates. VIII. The Isopoda. Amer. Natu- 
ralist 34:207-230. 

. 1 90 1. Key to the isopods of the Atlantic 

coast of North America with descriptions of 
new and little known species. Proc. U.S. Nat. 
Mus. 23:493-579. 

. 1904. Isopod crustaceans of the northwest 

coast of North America. Harriman Alaska 
Exped., Crustacea 10:213-230. [Reprinted: 
Richardson, H., 1904, Contributions to the 
natural history of the Isopoda, 2nd part, Proc. 
U.S. Nat. Mus. 27:657-681. ] 

. 1905a. A monograph on the isopods of 

North America. Bull. U.S. Natl. Mus. 54:1- 

. 1905/'. Isopods of the Alaska Salmon In- 
vestigation. Bull. U.S. Bur. Fisheries 24:209- 

. 1909. Isopods collected in the northwest 

Pacific by the U.S. Bureau of Fisheries steamer 
"Albatross" in 1906. Proc. U.S. Nat. Mus. 37: 


Rosen. D. E. 1978. Vicariant patterns and his- 
torical explanation in biogeography. Syst. Zool. 

Salemaa. H. 1979. Ecology of Idotca spp. (Iso- 
poda) in the northeast Baltic. Ophelia 18:133- 

Schminke. H. K. 1972. Evolution and homolo- 
gisierung der mandibeltypen der Bathynellacea 
(Crustacea, Malacostraca). Zcitschrift f. Zool. 
Systmatik Evolutionstbrshung 10:174-180. 

Schram, F. R. 1970. Isopod from the Pennsyl- 
vanian of Illinois. Science 169:854-855. 

. 1974. Paleo/oic Peracarida of North 

America. Fieldiana (Geology) 33:95-124. 

. 1978. Arthropods: a convergent phenom- 
enon. Fieldiana (Geology) 39(4):61-108. 

. 1981. On the classification of Eumala- 

costraca. J. Crustacean Biol. 1(1):1-10. 

Schultz. G. A. 1979. Aspects of the evolution and 
origin of the deep-sea isopod crustaceans. Sar- 
sia 64:77-83. 

Shackleton, N. J., and J. P. Kennett. 1975. Pa- 
leotemperature history of the Ceno/.oic and the 
initiation of Antarctic glaciation: oxygen and 
carbon isotope analyses in DSDP sites 277. 279 
and 281. Initial Rpts. Deep Sea Drilling Proj. 

Sheppard, E. M. 1957. Isopod Crustacea. Part II. 
Discovery Rpts. 29:141-198. 

Siewing, R. 1958. Anatomic und histologic von 
Thermosbaena mirabilis. Ein beitrag zur phy- 
logenie der reike Pancarida (Thermosbaena- 
cea). Abh. Math.-Naturw. KJ. Akad. Wisk. 
Mainz 7:197-270. 

. 1963. Studies in malacostracan mor- 
phology: results and problems. Pp. 85-103 m 
H. B. Whittington and W. D. I. Rolfe (eds.). 
Phylogeny and Evolution of Crustacea. Spec. 
Publ., Mus. Comparative Zool. Harvard Univ. 
Press, Cambridge. 

Simberloff. D.. K. L. Heck. E. D. McCoy, and E. 
F. Conner. 1981. There have been no statis- 
tical tests of cladistic biogeographic hypothe- 
ses. Pp. 40-63 in G. Nelson, and D. E. Rosen 
(eds.). Vicariance Biogeography: A Critique. 
Columbia Univ. Press, New York. 593 pp. 

Smith, A. G. and J. C. Briden. 1977. Mesozoic 
and Cenozoic Paleocontinental Maps. Cam- 
bridge Univ. Press, London. 63 pp. 

Stebbing. T. R. R. 1893. A History of Crustacea. 
Recent Malacostraca. D. Appleton and Co., 
New York. 466 pp. 

. 1905. Report on the Isopoda collected by 

Professor Herdman, at Ceylon, in 1902. Pp. 1- 
64 in W. A. Herdman, Report to the govern- 
ment of Ceylon on the pearl oyster fisheries of 
the Gulf of Manaar, with supplementary re- 
ports upon the marine biology of Ceylon by 
other naturalists. Part IV. Suppl. Rpt. No. 23. 
Royal Soc. London. 

Note added in proof: Kussakin (1982) recently synonymized west Pacific records of Cteantioides planicauda 
(as Cleantis planicauda) with his new Zenobiana rotunda Kussakin, 1982. This move restricts the genus 
Cleantioides to the tropical Pacific and Atlantic waters of the New World. (Kussakin. O.. 1982. Marine and 
brackish-water Isopoda of cold and temperate waters of the Northern Hemisphere. II. .Anthuridae. Microcer- 
beridae, Valvifera, Tyloidea. [in Russian] Opredeliteli po faune SSSR, Akad. nauk SSSR [Acad. Sci. USSR. 
Zool.]. No. 131. 

Steele, D. H. and V. J. Steele. 1975. Egg size and 
duration of embryonic development in Crus- 
tacea. Internat. Revue Ges. Hydrobiol. 60(5): 

Stock, J. H. 1981. L'origine geologique des lies 
des Indes Occidentales en relation avec la dis- 
persion de quelques Malacostraces stygo- 
biontes. Beobios No. 14(2):2 19-227. 

Sywula, T. 1964. A study on the taxonomy, ecol- 
ogy and the geographical distribution of species 
of genus Idotea Fabricius (Isopoda, Crustacea) 
in Polish Baltic. II. Ecological and zoogeo- 
graphical part. Bull. Soc. Amis. Sci. Lett. Poz- 
nan (D)4: 1 73-199. 

Tait, J. 1917. Experiments and observations on 
Crustacea. Part IV. Some structural features 
pertaining to (ilvptonotus. Proc. Roy. Soc. 
Edinburgh 37:246-303. 

Taylor, M. E., and R. M. Forester. 1979. Distri- 
butional model for marine isopod crustaceans 
and its bearing on early Paleozoic paleozoo- 
geography and continental drift. Bull. Geol. Soc. 
Amer., Pt. 1, 90:405-413. 

Wallerstein, B. R., and R. C. Brusca. 1982. Fish 
predation: a preliminary study of its role in the 
zoogeography and evolution of shallow water 
idoteid isopods (Crustacea: Isopoda: Idotei- 
dae). J. Biogeography 9:135-150. 

Watling, L. 1981. An alternative phylogeny of 
peracarid crustaceans. J. Crustacean Biol. 1(2): 

Watrous, L. E. and Q. D. Wheeler. 1981. The out- 
group comparison method of character anal- 
ysis. Syst. Zool. 30(1): 1-11. 

Wiley, E.O. 1981. Phylogenetics. The Theory and 
Practice of Phylogenetic Systematics. John Wi- 
ley and Sons, New York. 439 pp. 

Williams. W. D. 1970. A revision of North Amer- 
ican epigean species of Asellus (Crustacea: Is- 
opoda). Smithsonian Contrb. Zool. No. 49:1- 

Woodring, W. P. 1954. Caribbean land and sea 
through the ages. Bull. Geol. Soc. America 65: 

. 1966. The Panama land bridge as a sea 

barrier. Proc. Amer. Phil. Soc. 1 10:425-433. 

Zinsmeister, W. J. 1974. A new interpretation of 
thermally anomalous molluscan assemblages 
of the California Pleistocene. J. Paleontology 

. 1976. Biogeographical significance of the 

Late Mesozoic and Early Tertiary molluscan 
faunas of Seymour Island (Antarctic Peninsula) 
to the final breakup of Gondwanaland. Pp. 349- 
355 in J. Gray, and A. J. Boucot (eds.). His- 
torical Biogeography, Plate Tectonics, and the 
Changing Environment. Oregon State Univ. 
Press, Corvallis. 500 pp. 



Volume 20 Number 8 pp. 135-144 18 January 1984 

Rhamdia reddelli, new species, the first blind pimelodid catfish 
from Middle America, with a key to the Mexican species 

Robert Rush Miller 

.v / ^n /•« / /c HARVARD 

Museum oj Zoology ana Division of Biological Sciences, 

The University of Michigan. Ann Arbor. Michigan 48109 USA i_NllV 

Abstract. A new blind, depigmented catfish is described and illustrated from Cueva del Nacimiento 
del Rio San Antonio, Oaxaca, Mexico, and compared with its closest relative. Rhamdia laticauda. It 
resembles that species in having strong, retrorse serrae on the pectoral spine, and a shallowly notched 
caudal fin, but differs in the longer head, longer adipose fin, larger cephalic sensory pores, and longer 
and more numerous gill rakers (1 1-16 vs. 9-12). The karyotype (2n = 58) of the new species and a key 
to described Mexican species are given. A list of nominal and misidentified Mexican species is presented 
and R. laticauda. R. parryi. and R. guatcmalensis are illustrated. 

Resumen. Una nueva especie de bagre anoftalmo y depigmentado de la familia Pimelodidae se 
describe de la Cueva del Nacimiento del Rio San Antonio, Oaxaca, Mexico. Se distingue de Rhamdia 
laticauda. especie estrechamente relacionada a ella, por la cabeza mas grande y larga, la aleta adiposa 
mas larga, poros cefalicas mas grandes, y por la longitud y el numero de las branquispinas (11-16 vs. 
9-12). El numero de las cromosomas(2n = 58), clavesparaladeterminacion delosespeciesde Rhamdia. 
y illustraciones de R. laticauda. R. parryi, y R. guatcmalensis se presentan, y las especies describidas 
de Mexico se listan. 

Heresay reports of blind catfish from Mexico and Central America have persisted 
since before the turn of the century. As yet, however, the only described eyeless species 
from this region has been Prietella phreatophila Carranza (1954), from northeastern 
Mexico, a member of the Nearctic family Ictaluridae (6 genera, nearly 40 species). 
Troglobitic species apparently have evolved at least three times in this family (Lundberg 
1982; Reddell 1981:243-244, gives many references to Prietella). The Neotropical 
catfish family Pimelodidae is much larger (about 56 genera and 290 species according 
to Nelson 1984) but. thus far, only three blind species have been described (see Thines 
1955, for references). Two of these are from Sao Paulo, Brazil {Pimelodella kronei and 
Caecorhamdella brasiliensis), the third from Trinidad {Rhamdia urichi). A cave pop- 
ulation with the eye variably reduced was described recently from Belize as Rhamdia 
laticauda lyphla (Greenfield et al. 1983). 

For many years it was common practice to assign cavernicolous fishes to distinct 
genera even though they typically differed from their epigean relatives only in lacking 
eyes and being depigmented. That viewpoint has changed markedly in recent years (see 
discussions by Roberts and Stewart 1 976, and Banister and Bunni 1 980) and a number 
of blind fishes originally placed in monotypic genera have been reassigned to their more 
widespread surface relatives. In proposing the genus Caecorhamdia for Rhai)idia urichi, 
Norman (1926) wrote that his genus differed from Rhamdia only in lacking eyes and 
that C. urichi was "almost identical" with Rhamdia quelen, the type species o{ Rhamdia. 
Mees (1974:152, 160) agreed, placed Caecorhamdia in synonymy with Rhamdia. and 
designated Norman's species as RhaDidia quelen urichi. Haseman (1911:325) stated 
that Typhlobagrus kronei is indistinguishable from Pimelodella lateristrigata, except 
for the loss of sight, and recommended that the cave form be relegated to subspecific 


Figure 1. Rhamdia reddelli. A, Holotype (5?), UMMZ 21 1 164, 98.5 mm SL. 

Catfishes have anatomical, physiological, and behavioral characteristics that prea- 
dapt them to life in darkness (e.g., well developed organs of taste and touch, nocturnal 
activity, crevice-seeking habits). It is not surprising, therefore, that about 40 percent 
(17 of 38 species) of the blind fishes that inhabit fresh water are siluroids. Among the 
five genera of pimelodids inhabiting Middle America, only Rhamdia is widely distrib- 
uted and evolutionarily successful (Bailey and Miller, 1979). The Mexican and Central 
American representatives of this genus are under review by Reeve M. Bailey and myself; 
some of our conclusions receive advance notice in this paper. 

The eyes of Rhamdia are normally small and of secondary importance in their 
life. Field observations in Honduras by Carr and Giovannoli (1950) oi Rhamdia bra- 
chycephala indicate that this species (a synonym of R. cabrerai—see below) is ''exclu- 
sively cavernicolous and thigmotactic" in its swift-water habitat. 

The troglobitic species described below differs from its epigean relatives in many 
features other than depigmentation and loss of eyes. It may be known as: 

Rhamdia reddelli new species 
Figures 1-3 

Synonymy.— Rhamdia new species— LeGrande, 1981:42 (chromosome and arm 
numbers, based on UMMZ 199016). Reddell, 1981:244-245 (mentioned; type locality 

Holotype. — UMMZ 21 1 164, a male? 98.5 mm SL, Cueva del Nacimiento del Rio 
San Antonio, ca. 9 km SW of Acatlan, Oaxaca, on Atlantic slope of eastern Mexico; 
collected by James R. Reddell and Andy G. Grubbs, 2 January 1977. 

Paratypes.—W\ specimens are from same locality as holotype. UMMZ 199016 (2 
specimens: 39 and 70 mm), A. G. Grubbs, M. Cossey, and T. Byrd, 8 January 1976, 
shipped alive to Ann Arbor (larger individual karyotyped); UMMZ 21 1 165 (6 speci- 
mens: 5 1 .5-94.2 mm), taken with the holotype; UMMZ 2 1 1 1 66 (77. 1 mm), R. Mitchell 
and L. Faulkenberry, 7 January 1977. AMNH 38216 (98.2 mm), J. Reddell, D. and 
M. H. McKenzie, S. Murphy, 26 December 1972; AMNH 38217 (4 specimens: 36.7- 
90.5 mm), same collectors and date; AMNH 38218 (2 specimens: 68.5, 69.5 mm), 
same collectors. 9 March 1973. 

Diagnosis. — A species of Rhamdia with a broad, moderately depressed skull in 
adult, long head, very weak and short occipital process, almost no pigmentation, and 
lacking eyes (a tiny eye spot occurs in a 37-mm specimen). Related to R. laticaiida 
which it resembles in the strong, retrorse serrae on the posterior edge of the pectoral 
spine and in the shallowly notched caudal fin. From R. laticauda and its closest relatives 
{see below) it differs in having: (1) a much longer head and adipose fin, (2) cavernous 
sensory head pores (especially on chin), and (3) longer and more numerous gill rakers 


Figure 2. Rhamdia reddelli. Lateral (A), ventral (B). and dorsal (C) views of paratype, AMNH 38216. 98.2 
mm SL. Arrows indicate tips of pectoral spines. The fork of the caudal fin is too deep as drawn. 

(11-16 VS. 9-12 on first arch). Head enters SL 3.25-3.75 times (vs. 4.0-5.5) and the 
depressed dorsal fin overlaps the adipose fin. 

Description. — Body form and color pattern are indicated in Figures 1-2. Propor- 
tional measurements are presented in Table 1. Meristic data (based on 16 fish) follow. 
Gill rakers were counted on the first (right) arch, with numbers for upper and lower 
limbs recorded separately (raker at angle included in lower-limb count). Vertebral 
counts are post-Weberian. with separation of precaudal and caudal counts where pos- 
sible (5 vertebrae comprise the Weberian complex). Dorsal fin invariably 1,6. the spine 
soft and flexible as typical oi^ Rhamdia: anal rays 13 or 14 (anterior rudiments difiicult 
to see); pectoral rays 1,10 or 1,1 1, usually 1,10 (22 of 30 counts); pelvic rays invariably 
6; principal caudal rays 17-19 (16-18 branched). Gill rakers long, slender, 3 + 8 to 4+ 12, 
total 11(2), 12(0), 13(5), 14(6), 15(1). 16(1). Vertebrae: precaudal, 7 or 8, caudal. 29- 
32, total 37-39. The number of posterior serrae on the pectoral spine varies with size, 
from 6-6 in a 36.7-mm SL specimen to 15-14 in a 90.5-mm SL specimen. In the 
larger fish the serrae are triangular, with very broad bases, and, except proximally. there 
is no gap between individual serrae as in Rhamdia laticauda and closest relatives {R. 
parryi. R. salvini, R. cabrcrai^). The pectoral spine is gently curved in adults but in 
specimens less than 70 mm SL it is straight and there are gaps between the individual 
serrae. There are no serrae on the anterior edge of the spine, which is essentialh smooth. 

The maxillary barbel is generally longer than in R. laticauda and much longer than 
in R. parryi, R. salvini. or R. cahrerai, but it is shorter than in R. guatemalensis which 
belongs to a diflferent species group. It may extend backward almost as far as the tip 
of the depressed dorsal fin and well beyond the origin of the adipose fin. but in some 

Meek (1906) named this fish for Senor Cabrera but spelled the patronym cabrerae. It is here corrected. 


8S ^ ^ SI SS ^ « 





1^ ^ ^ u ^ ft A. 

^ m^ ^ J^ {^ #^ 

FiGi RE 3. Somatic chromosomes at metaphase of female paratype, UMMZ 199016: 2n = 58. FN = 100±4 
(LeGrande, 1981:42). 

it does not reach beyond the middle of the depressed pectoral fin. The insertion of the 
pelvic fin lies before the end of the dorsal-fin base. The long adipose fin is well developed, 
especially posteriorly. The occipital process is very weak and short, extending less than 
one-fourth the distance to the dorsal-fin origin. 

Alive in its natural habitat, Rhamdia reddelli is virtually colorless and without 
visible pigment. In ethyl alcohol (formalin-fixed), the holotype (Fig. 1 ) has fine, scattered 
flecks of pigment on the top and sides of the head, along the back and upper sides, and 
in a narrow band along the lateral line. The lower sides and entire venter are immaculate, 
as are all the fins except the caudal which has the interradial membranes largely dusky. 
The similar-sized paratype (Fig. 2) also essentially lacks pigment except for fine flecks 
along and below the base of the adipose fin and some duskiness on the interradial 
membranes of the caudal fin. Other adults (between 69 and 9 1 mm SL), except for one 
mentioned below, resemble either the holotype or the above-described paratype, or the 
caudal fin may be immaculate. An 81.5-mm specimen (UMMZ 21 1 165) is more pig- 
mented, with fine flecks extending downward to a line just above the bases of the 
pectoral, pelvic, and anal fins and with fine pigment grading onto the ventral surface 
of the caudal peduncle; the venter is otherwise immaculate. Some juveniles (3 or 4 in 
UMMZ 21 1 165) are pigmented as fully as the holotype, whereas others virtually lack 
pigment. When kept alive for a month or more in a lighted place, fine melanophores 
spread over much of the body, covering the venter posterior to the anus and encroaching 
anteriorly onto the abdomen, with small ones developing along the rays of the paired 


Table 1 . Proportional measurements (in permillage of SL) of 1 2 specimens (juv.-ad.) of the types of Rhamdia 
reddelti. The data for the holotype are included in the summary. Figures in parentheses are number of 
specimens when fewer than 12. 





Standard length, mm 

Body depth 

Predorsal length 

Preanal length 

Anal origin to caudal base 

Caudal-peduncle length 

Caudal-peduncle depth 

Head length 

Head depth 

Head width (11) 

Snout length 

Mouth width 

Interorbital width 

Maxillary-barbel length (9) 

Outer mental barbel length (11) 

Inner mental barbel length (11) 

Adipose-fin length 

Adipose-fin maximum height 

Anal-fin basal length 

Pectoral-fin length (11) 

Pectoral-spine length 

Caudal-fin length (9) 

Caudal fin, to notch (8) 

Caudal fin, shortest ray length (8) 









































































fins and the anal fin as well as on the interradial membranes of the dorsal fin (e.g., the 
69-mm SL specimen, UMMZ 199016, kept alive over six months). The one individual 
with a tiny pigmented pupil (AMNH 38217, 36.7 mm SL) appears to lack pigment. 

From the recently described Lake Nicaraguan species Rhamdia hiigiana Villa 
(1977). the new species differs markedly in the much deeper body, shorter pectoral 
spine and fin. longer head, shorter maxillary barbel, shallower caudal-fin notch, and 
rounded rather than pointed caudal-fin lobes. It also has fewer post- Weberian vertebrae 
(37-39 vs. 39-42 in R. luigiana). Rhamdia reddelli is compared with other Mexican 
species of Rhamdia in the Key. 

The karyotype (Fig. 3), kindly prepared by William H. LeGrande, shows a diploid 
number of 58 chromosomes and an arm number {¥N) of 100±4. This diploid number 
may be the ancestral condition for pimelodids, as it evidently is for ictalurids (LeGrande 
1981), but since the karyotypes of only seven species of pimelodids have been published 
(2n = 46, 56, 58, 62— see LeGrande 1981:42) this tentative conclusion must await 
further karyological studies of this family. 

Habitat and associates.— Jho: cave is at Canada San Antonio, approximately 9 km 
SW of Acatlan, at an elevation of 100 m. The stream flowing from below the cave 
entrance is the primary source of the Rio San Antonio and drains south into Presa 
Miguel Aleman, a dammed portion of the Rio Tonto that is tributary to Rio Papaloapan. 
The main passage of the cave extends for about 120 m to a deep lake containing many 
blind catfish and crayfish. Beyond this lake a shallow stream extends for 350 m in a 
passage 10 to 30 m wide and up to 1 1 m high. Several major side passages that contain 
secondary streams occur throughout the cave and bring the total length of the cave to 
about 4.5 km. The main stream floor is generally of sand and gravel with areas of 
flowstone and bedrock. Both air and water temperatures were 23. 5°C. The catfish usually 
occurred in the deeper ponded portions of the streams, especially in areas over which 
bats roosted. 

A rich invertebrate cave fauna is associated with the catfish. Four species of trog- 
lobitic crustaceans inhabit the cave: Potamalpheops stygicola Hobbs (Decapoda: 


Alpheidae), Macrobrachiu!)i villalobosi Hobbs (Decapoda: Palaemonidae). Procam- 
banis (Austrocaniharus) oaxacae rcddclli Hobbs (Decapoda: Cambaridae), and Speleo- 
mysis olivae Bowman (Mysidacea: Lepidomysidae). All have since been collected in 
other caves in the vicinity of Acatlan. A second species of mysid, Antromy»is (Antro- 
mysis) reddelli Bowman, has been collected from a nearby cave and can be expected 
to occur in Cueva del Nacimiento del Rio San Antonio. A specimen of the alpheid 
shrimp Potanialpheops stygicola was disgorged by a catfish upon preservation. The 
rarity of shrimps and mysids in pools containing catfish is doubtless related to predation 
by the fish on the crustaceans. The cave is also inhabited by a possibly troglobitic clam, 
which is abundant in various parts of the cave but awaits study. 

The terrestrial fauna is extremely abundant and includes troglobitic trichoniscid 
isopods, nicoletiid thysanurans, millipeds, spiders, and opilionids. 

Etymology. — \ am pleased to name this distinctive species for James R. Reddell, 
who donated all of the type specimens and has pioneered in exploring caves in Latin 

Nominal or Misidentified Species Referred 
TO Mexican Catfishes of the Genus Rhamdia 

Piinelodiis laticaudus Meckel in Kner. 1858 (Abtheilung Sitzber. Akad. Wiss. Wien, 26: 
420). Type locality (on label in jar): "Rio Xamapa, Mexiko." Oldest available 
name for a Middle American Rhamdia. A valid species. Three syntypes, Vienna 
Museum 50554 (166, 171, 203 mm SL), examined. 

Pi melodus guatema/ensis Gunthev, 1864 (Cat. Fish. British Mus., 5:122). Type locality: 
Huamuchal, on Pacific coastal plain, Guatemala. A valid species. 

Pimelodus godmani Giinther, 1 864 (ibid.: 1 24). Type locality: Guatemala (Rio Motagua, 
lower Vera Paz) and Mexico. Species illustrated by Regan (Biol. Centrali-Ameri- 
cana, 8:pl. 21, fig. 1). A synonym o^ Rhamdia guatemalensis (Miller 1966:787). 

Pimelodus petenensis Giinther, 1864 (ibid.: 126). Type locality: Lake Peten, Guatemala. 
Listed for Mexico by Alvarez 1 950 (Sec. de Marina, Dir. Gen. Pesca e Ind. Conexas, 
Mexico:35) with the remark ''probably only in Guatemala." Illustrated by Regan 
(op. cit.:pl. 22, fig. 1). A subspecies of/?, guatemalensis (Hubbs 1938:266). 

Pimelodus hypselurus Giinther, 1864 (ibid.: 126-1 27). Type locality (on label in jar): 
Orizaba [but listed as Cordova in cat. book], Mexico. Holotype, BMNH 1858- 
1 1.22.32 (103.5 mm SL), examined by R. M. Bailey; illustrated by Regan (op. cit.: 
pi. 21, fig. 3). A synonym of R. laticauda. 

Pimelodus motaguensis Giinther, 1864 (ibid.: 127). Type locality: Rio Motagua, Gua- 
temala. Holotype illustrated by Regan (op. cit.:pl. 20, fig. 1). A synonym oi R. 
laticauda (see comment by Miller 1976:4). 

Pi?nelodus brachypterus Cope. 1866 (Trans. Amer. Philos. Soc, 13:404). Type locality: 
Orizaba, Mexico. Holotype, ANSP 16471 (147 mm SL), examined. A synonym 
of R. laticauda. 

Rhamdia parryi Eigenmann and Eigenmann, 1888 (Proc. Calif Acad. Sci., ser. 2, vol. 
1:130). Type locality: Rio Zanaleneo [=Sanatenco], near Tonala, on Pacific slope 
of Chiapas, Mexico. Five syntypes, MCZ 27273 (77-88 mm SL), examined. A 
valid species (called R. hypselura by Miller 1966:787). 

Rhamdia oaxacae Meek, 1902 (Field Col. Mus. Publ. 65:74, pi. 14). Type locality: Rio 
Quiotepec at Cuicatlan, Oaxaca, Mexico, in Rio Papaloapan basin. A synonym of 
R. guatemalensis (Regan op. cit.: 132). 

Rhamdia depressa Barbour and Cole, 1906 (Bull. Mus. Comp. Zool. 50(5): 155, pi. 1). 
Type locality: Ikil [=Ixil] Cenote near Chichen-Itza, Yucatan, Mexico. A subspecies 
of R. guatemalensis (Huhbs 1936:194). 

Rhamdia sacrificii Barbour and Cole, 1906 (ibid.: 156). Type locality: Sacrificial cenote 
near Chichen-Itza, Yucatan, Mexico. A synonym of R. guatemalensis (Hubbs 1 936: 

Pimelodus houcardi Regan, 1907 (Ann. Mag. Nat. Hist., ser. 7, vol. 19:258). Type 





-^ ^ 










 -■■■in l.iii^i 1 "T " 



FicH'RE 4. Three species of Rhanidia from Mexico. A, 7?. guatemalensis. UMMZ 183901 (126.2 mm). 
Nacimiento del Rio Cosalapa, 2.6 km SE of Estacion Tezonapa, Veracruz; B. R. laticauda. UMMZ 196674 
(68.8 mm), tributan, to Rio Metlac. Fortin. Veracruz; C, R. parryi. UMMZ 184739 (68.0 mm), headwaters 
of Rio Tapanatepec, at Hwy 190 bridge E of Tapanalepec, Oaxaca. 

locality: Yucatan. Mexico. A synonym of R. guatemalensis depressa Barbour and 
Cole (Hubbs 1936:193, 195); illustrated by Regan (op. cit.:pl. 20, fig. 3). 
Pimehdiis brachycephalus Regan, 1 907 (op. cit.:258). Type locality: Rio Nacasil, Pacific 
slope of Guatemala. Recorded tentatively from Mexico by Alvarez 1950 (op. cit.: 
37) with the remark "probably only in Guatemala." Illustrated by Regan (op. cit.: 
pi. 22, fig. 2). A synonym of R. cahrerai (type examined by R. M. Bailey). Later. 
Alvarez (1970:77) listed R. hrachycephala from "southeastern Mexico near the 


Guatemalan frontier," but this represents a misidentification since R. cabrerai is 
known on the Atlantic slope of Guatemala only from the upper Rio Motagua. 

Key to Mexican Species of Rhamdia- » 

R. laticauda and R. parryi are commonly sympatric with R. guatemalensis. 

la. Anterior and posterior edges of pectoral spine with small, numerous serrae of 
subequal length, developed about equally or those on posterior edge somewhat 
stronger (especially in older fish); caudal fin deeply notched for at least two- 
thirds the distance from tips of caudal lobes to base of mid-caudal rays. 
(Maxillary barbel long, typically extending well beyond origin of adipose fin; 
head long, 3.5-3.8 in SL; adipose fin long, ca. one-third SL; occipital process 
long, extending nearly halfway or more to dorsal origin.) Atlantic and Pacific 
lowlands from just NW of Veracruz City on Atlantic slope and Rio Tehuan- 
tepec basin on Pacific versant southward to Panama (if R. wagneri is a syn- 
onym— 56'e' Hubbs 1936:181); typically in pools 

Rhamdia guatemalensis (Fig. 4) 

b. Pectoral spine with strong, retrorse to nearly straight serrae only on posterior 
edge (anterior edge smooth or roughened); caudal fin weakly notched, to no 
more than half distance from tips of caudal lobes to base of mid-caudal rays. 
Atlantic and Pacific versants\ cavernicolous and in rocky streams of piedmont 

slopes and foothills 2 

2a. Skull depressed; head long, 3.25-3.75 in SL; blind and depigmented; adipose 
fin well developed, overlapped by depressed dorsal fin. Cueva del Nacimiento 
del Rio San Antonio, Oaxaca; cavernicolous Rhamdia reddelli (Figs. 1-2) 

b. Skull domed; head short, 4.0-5.5 in SL; eyes and pigment well developed; 

adipose fin short, not (or rarely) overlapped by depressed dorsal fin 3 

3a. A prominent, dark lateral stripe on midside, from behind head to base of 
caudal fin, becoming broader posteriorly; post-Weberian vertebrae fewer, 35- 
38, usually 36 or 37 (98%). Pacific slope of Oaxaca and Chiapas southeastward 

into Guatemala (to Dpto. de Santa Rosa); on rocky riffles 

Rhamdia parryi (Fig. 4) 

b. Side of body without a conspicuous dark stripe; post-Weberian vertebrae more 
numerous, 37-41, usually 38-40 (91%). Atlantic slope from Rio Jamapa, 
Veracruz, southeastward to western Honduras; on rocky riffles and in current 
of streams Rhamdia laticauda (Fig. 4) 


Robert W. Mitchell called my attention to the existence of the new species. James 
R. Reddell provided the information from which the account of the habitat and as- 
sociates was written. 

I am grateful to the following for information, specimens, radiographs, and other 
assistance, and for loans of important material needed for the completion of this paper: 
P. H. Greenwood, British Museum of Natural History (BMNH), Paul Kahsbauer, 
Vienna Museum, Donn E. Rosen, American Museum of Natural History (AMNH), 
James E. and Eugenia B. Bohlke, Academy of Natural Sciences of Philadelphia (ANSP), 
Karel Liem and William L. Fink, Museum of Comparative Zoology at Harvard (MCZ), 
Robert H. Gibbs, Stanley H. Weitzman, and Susan L. Jewett, United States National 
Museum of Natural History (USNM), Reeve M. Bailey, Charles E. Dawson, Andy G. 
Grubbs, William H. LeGrande, Doyle Mosier, Marcia K. Dorsey, and Alexandra Snyder 
Creighton. Louis P. Martonyi (Fig. 4B), Edward C. Theriot (Figs. 1, 4C), and William 
M. Pelletier (Fig. 4A) are responsible for the photographs, Anita Cleary for the drawings 
(Fig. 2). Margaret L. Van Bolt mounted Figure 2 and added heavier arrows. Mexican 

Much of this key was derived from information provided by R. M. Bailey. 


officials permitted me to conduct the field work, and the National Science Foundation 
supported field and laboratory work (most recently as grant DEB 80-02017). Three 
anonymous reviewers improved the manuscript. 

Literature Cited 

Alvarez. J. 1970. Feces mexicanos (claves). Inst. 
Nac. Inv. Biol. Pesq., Ser. Inv. Pesq.. Estudio 
No. 1, Mexico. 166 pp. 

Bailey. R. M., and R. R. Miller. 1979. Pimelodid 
catfishes (genus Rhamdia) from northern Mid- 
dle America. .Abst. 59th .Ann. Mtg. Amer. Soc. 
Ich. & Herp.. Orono. Maine. 1 p. 

Banister. K. E.. and M. K. Bunm. 1980. A new 
blind cyprinid fish from Iraq. Bull. Brit. Mus. 
Nat. Hist. (Zool.) 38(3): 15 1-158. 

Carr. A. F. Jr.. and L. Giovannoli. 1950. The 
fishes of the Choluteca drainage of southern 
Honduras. Occ. Pap. Mus. Zool. Univ. Mich. 

Carranza. J. 1954. Descripcion del primer bagre 
anoftalmo y depigmentado encontrado en aguas 
mexicanas. Ciencia 1 4(7-8): 129-1 36. 

Greenfield. D. W.. T. A. Greenfield, and R. L. 
Woods. 1983. A new subspecies of cave- 
dwelling pimelodid catfish, Rhamdia laticauda 
tvphla. from Belize. Central America. Brenesia 

Haseman, J. D. 1911. Descriptions of some new 
species of fishes and miscellaneous notes on 
others obtained during the expedition of the 
Carnegie Museum to central South America. 
Ann. Carnegie Mus. 7(3-4):3 15-328. 

Hubbs, C. L. 1936. Fishes of the Yucatan Pen- 
insula. Carnegie Inst. Wash. Publ. 457:157- 

. 1938. Fishes from the caves of Yucatan. 

Carnegie Inst. Wash. Publ. 491:261-295. 

LeGrande. W. H. 1981. Chromosomal evolution 
in North American catfishes (Siluriformes: Ic- 
taluridae) with particular emphasis on the 
madtoms, Notunis. Copeia 1981(l):33-52. 

Lundberg. J. G. 1982. The comparative anatomy 

of the toothless blindcat, Trogloglams paller- 

soni Eigenmann, with a phylogenetic analysis 

of the ictalurid catfishes. Misc. Publ. Mus. Zool. 

Univ. Mich. 163:1-85. 
Meek, S. E. 1906. Description ofthree new species 

of fishes from Middle America. Field Col. Mus. 

Publ. 1 16, Zool. Ser. 7(3):93-95. 
Mees, G. F. 1974. The Auchenipteridae and Pi- 

melodidae of Suriname (Pisces, Nematogna- 

thi). Zoolog. Verhand. 132:1-256 + pis. 1-15. 
Miller, R. R. 1966. Geographical distribution of 

Central American freshwater fishes. Copeia 

. 1976. An evaluation of Seth E. Meek's 

contributions to Mexican ichthyology. Field- 

lana. Zool. 69(1): 1-31. 
Nelson, J. S. 1984. Fishes of the World. 2nd ed. 

John Wiley & Sons, N. Y. 416 pp. 
Norman, J. R. 1926. A new blind catfish from 

Trinidad, with a list of the blind cave-fishes. 

Ann. Mag. Nat. Hist. ser. 9, vol. 18:324-331. 
Reddell, J. R. 1981. A review of the cavernicole 

fauna of Mexico, Guatemala, and Belize. Texas 

Mem. Mus., Bull. 27:1-327. 
Roberts, T. R., and D. J. Stewart. 1976. An eco- 
logical and systematic survey of fishes in the 

rapids of the lower Zaire or Congo River. Bull. 

Mus. Comp. Zool.. Harvard Univ. 147(6):239- 

Thines. G. 1955. Les Poissons aveugles (1): ori- 

gine, taxonomie. repartition, geographique. 

comportement. Ann. Soc. Roy. Zool. Belgique 

Villa, J. 1977. A new species of pimelodid catfish 

of the genus Rhamdia from Nicaragua, Central 

America. Brenesia 12/13:133-142. 





[OCT 1 1984 

Volume 20 Number 9 pp. 145-150 20 June 1984 

A complete specimen of Peachella brevispina Palmer— an unusual 
olenellid trilobite (Arthropoda: Olenellida) from the 
lower Cambrian of California 

James H. Stitt 

Department of Geology, University of Missouri-Columbia, Columbia, MO 65211 USA 

R. L. Clark 

Department of Geology, Paleontology Section. San Diego Natural History Museum, 
P.O. Box 1390. San Diego, C A 92112 USA 

Abstract. The thorax and possible pygidium of Peachella brevispina are described for the first 
time from a recently collected complete specimen, the first recovered for this genus. Unusual features 
include the large macropleural third segment of the prothorax that distorts the adjacent pleurae, the 
rounded distal extremeties of the other prothoracic segments, the rather gradual transition from the 
prothorax to the opisthothorax, and the large, triangular pygidium. 


Olenellid trilobites are found in Lower Cambrian strata in various parts of the 
world, and numerous specimens have been collected from localities in western North 
America. Olenellids are broad, flat, medium to large-sized trilobites characterized by 
a large semicircular cephalon, prominent ocular lobes, numerous thoracic segments 
that terminate laterally in spines of various lengths, and a small pygidium. The absence 
of specialized morphologic features on the thorax that were used for enrollment in 
other trilobites (Bergstrom 1973:17) indicates that trilobites of the Family Olenellidae 
were unable to enroll (Harrington 1959:102), a protective feature that developed in 
other Lower Cambrian (Rasetti 1948:17-18, pi. 4, figs. 22-24) and later trilobites. 

Initial taxonomic efforts on olenellid trilobites resulted in the recognition of a 
number of closely related genera (Walcott 1910) and species (Resser and Howell 1938). 
More recent work (Fritz 1972, Palmer and Halley 1979) has resulted in recognition of 
fewer genera, because the criteria originally used to discriminate certain genera inter- 
grade among various species. Added to this problem is the fact that olenellids frequently 
occur in the same beds of rock in pairs of closely related taxa that have been variously 
interpreted as paired species or dimorphs (see Palmer and Halley 1974:66-67 for a 
recent discussion of these taxonomic problems). 

The complete specimen of Peachella brevispina Palmer described in this paper was 
collected by R. L. Clark from the lower part of the Carrara Formation at Emigrant 
Pass in the Nopah Range, Inyo County, California (Fig. 1). Palmer and Halley (1979: 
13, 75; fig. 1) reported that P. brevispina is moderately rare in the Thimble Limestone 
Member of the Carrara in the nearby Dublin Hills and at Eagle Mountain. Mount 
(1980:78-80; fig. 1) described the Emigrant Pass locality in the Nopah Range, including 
a detailed columnar section and range chart in which he reported P. brevispina from 
a thin limestone bed near the top of an unnamed lower member of the Carrara. Mount's 
columnar section supports Palmer and Halley's observation (p. 9, 13; figs. 6A, 1 1) that 
the Thimble Limestone Member of the Carrara cannot be recognized in the Nopah 
Range because the Thimble grades southeastward into shale. 


Peachella is an unusual genus of olenellid trilobites that was known only from a 
small number of cephala before Clark's discovery of the complete specimen described 
in this paper. The glabella, ocular lobes and cephalic border are fainter and less well 
defined than usual for an olenellid. The most unusual feature of the ceph^on is the 
genal spines, which are normally long and pointed in other olenellids but in species of 
Peachella are short, wide and have broadly rounded tips. First described by Walcott 
in 1910, this scarce genus included only the type species Peachella iddingsi (Walcott) 
until 1979, when Palmer described a second species, P. brevispina, from the Thimble 
Limestone. Clark's discovery of a complete specimen, which he donated to the San 
Diego Natural History Museum (SDSNH locality no. 3169), allows the description of 
this unusual trilobite to be completed. 

Morphologic terms used in the following description are defined in Harrington 
(1959). Suprageneric classification follows Bergstrom (1973) and Palmer and Halley 

Systematic Paleontology 

Phylum Arthropoda Siebold and Stannius, 1 845 

Class Trilobita Walch, 1771 

Order Olenellida Resser, 1938 

Family Olenellidae Vogdes, 1893 

Genus Peachella Walcott, 1910 

Peachella brevispina Palmer, 1979 

Figures 2, 3 

Peachella brevispina PALMER in Palmer and Halley (1979:75, pi. 5, figs 1-3). 

Material.— 1 complete decalcified specimen (SDSNH 24548) preserved as internal 
and external molds; specimen slightly deformed, especially the cephalon and pygidium. 

Description. — 'LtngXh of specimen 33 mm (excluding macropleural spines). Ceph- 
alon semicircular in outline with short, paddle-like genal spines. Anterior end of ce- 
phalon poorly preserved. Glabella prominent, elongate, extends close to anterior edge 
of cephalon, set off by rather shallow axial furrows. No lateral glabellar or occipital 
furrows visible. Occipital ring not differentiated. Ocular lobes poorly preserved, ap- 
parently short, arcuate and located close to the glabella. Anterior border and border 
furrow not preserved. Lateral border narrow opposite anterior end of glabella, gradually 
widens posteriorly. Lateral border furrow shallow, curved slightly adaxially at anterior 
end of genal spine, crossing adaxial base of genal spine and continuing to posterior 
margin of cephalon. Posterior border furrow not visible. Characteristic genal spines 
short, broad, moderately inflated, posterolaterally directed; spine termination broadly 

Prothorax of 1 5 segments. Axis prominent, convex, width (trans.) approximately 
one-fourth of prothoracic width. Most articulating half-rings visible on outstretched 
specimen. Axial furrows shallow. Axial ring and transverse furrow of first prothoracic 
segment slope steeply anteriorly, articulating half-ring apparently still connected to 
posterior ventral edge of cephalon. Axial ring of second prothoracic segment separated 
from narrow articulating half-ring by prominent, wide, deep transverse furrow that 
curves slightly anteriorly abaxially and intersects the axial furrows. Remaining axial 
rings separated from articulating half-rings by faint, narrow transverse furrow that 
curves anteriorly abaxially and terminates near anterior edge of ring before reaching 
axial furrows. This furrow deepest on axial rings of prothoracic segments 12-14, be- 
coming progressively fainter on anterior segments. Small axial node present on posterior 
edge of axial rings of prothoracic segments 10-14; node most prominent on segment 
14, progressively diminishing in size anteriorly. Prothoracic segment 15 broken at axis, 
undoubtedly bears axial spine whose mold can be seen entering the surrounding rock 
just posterior of the opisthothorax. 

Pleurae of prothorax variable in appearance. Pleurae of first two prothoracic seg- 


Figure 1. Map showing location of Emigrant Pass locality where described specimen was collected. 

ments not well preserved, appearing flat with no pleural furrows and rounded distal 
extremities. Faint transverse ridge(s) present on left pleurae of segments 1 and 2 not 
present on right pleurae, and interpreted to be result of deformation of specimen. Pleurae 
of segment 3 expand rapidly abaxially, especially along posterior edge, becoming broad- 
ly oval in shape and approximately seven times as long (exsag.) as the pleurae of any 
other segment. Pleurae of segment 3 reach maximum length (exsag.) near distal margins 
of thorax, pleurae narrow abaxial of maximum length to form large, long macropleural 
spine that curves posteriorly and extends well beyond the posterior end of the specimen. 
Unusual ovoid shape of pleurae of segment 3 distorts shape of pleurae of segments 2 
and 4-6, forcing these pleurae to compress abaxially and to taper distally (to rounded 
extremeties) in order to accommodate wider pleurae of segment 3 without leaving gaps 
or overlaps between pleurae. Pleurae of prothoracic segments 7-14 rather plain, nearly 
flat, with rounded distal extremeties. All pleurae posterior of third pleurae divided into 
narrow anterior and posterior bands by broad, shallow pleural furrow. Pleural furrow 
narrower and deeper in segments 12-14. Pleurae of prothoracic segments 10-14 bend 
gently to the posterior abaxially, with posterior bending greatest toward posterior end 
of prothorax. 

Opisthothorax consists of eight or nine segments. Axis only faintly defined by very 
shallow axial furrows. Pleurae simple, nearly flat, apparently with no pleural furrows 
and rounded distal extremeties. 




Figure 2. Stereophotographs of internal mold of specimen of Peachella brevispina Palmer (SDSNH 24548) 
from lower part of Carrara Formation, Emigrant Pass, California; a. entire specimen, x2; b. enlargement of 
thorax and pygidium, x4. 








if ' 


Figure 3. Stereophotographs ( x 1 6) of segments 1 3-1 5 of the prothorax, the opisthothorax, and the possible 
pygidium (p) of Peachella brevispina Palmer. The pygidium is poorly preserved and twisted underneath and 
to the right so that only the axis and right pleural region are visible. Note also that the axial ring of prothoracic 
segment 1 5 is broken. The base of the spine that extended upward and backward from this segment is 
preserved on the counterpart to this specimen, and the mold of this spine (s) enters the rock to the left of 
the possible pygidium. 

Pygidium possibly present, although twisted to right and only partly preserved on 
internal and external molds. Overall shape broadly triangular, with convex axis tapering 
posteriorly and extending almost to pygidial margin. Pleural area smooth, slightly 
convex, with no border or border furrow. 

Remarks. — Tht cephalon of Peachella brevispina is characterized by its faintly 
defined glabella and border furrow, short ocular lobes, and short, moderately inflated 
paddle-like genal spines. The cephala illustrated by Palmer and Halley (1979, pi. 5, 
figs. 1, 2) are much better preserved than the cephalon on this complete specimen. 

The most unusual features of the prothorax are the pleurae of the third segment, 
which expand rapidly away from the axial furrows and become broadly oval in shape, 
crowding and distorting the adjacent pleurae. Other unusual features include the round- 
ed distal extremeties of the prothoracic pleurae; on most olenellid trilobites. the pleurae 
terminate laterally in spines. The prothoracic segments diminish in size posteriorly and 
somewhat grade into the segments of the opisthothorax. although the junction between 
these two parts of the thorax is not perfectly preserved. Palmer (in Palmer and Halley 
1979:73) described a partly preserved thorax of Olenelliis multinodus (pi. 4, figs. 7, 8) 
that has enlarged pleurae on the third prothoracic segment and a gradual transition 
from the pleurae of the prothorax to the pleurae of the opisthothorax. He suggests that 
these features might merit placing O. multinodus in a new genus (possibly with Olenellus 
arcuatus) if additional specimens prove that these features are characteristic of the 
species. These two species also have short ocular lobes. Similar features are present on 
P. brevispina, which has in addition the unusual and generically distinctive paddle- 
shaped genal spines not present on O. multinodus or O. arcuatus. This mix of similar 


unusual features on species that clearly belong in different genera illustrates once again 
some of the problems in generic level taxonomy in olenellids. 

If indeed the pygidium of P. brevispina is preserved in a twisted position at the 
posterior end of the opisthothorax, then P. brevispina also has an unusuaLpygidium 
for an olenellid to add to its other peculiarities. Olenellid pygidia (when preserved) are 
very small, short trapezoidal plates attached to the end of the narrow opisthothorax. 
The apparent pygidium of P. brevispina is relatively large, triangular in shape, and has 
broad, smooth pleural areas, a combination which makes the pygidium of this species 
unique among olenellids. 


The senior author would like to thank Frederick Schram of the San Diego Natural 
History Museum for suggesting the project. Three anonymous reviewers are thanked 
for their helpful comments and suggestions. Connie Egerdahl is thanked for carefully 
typing the manuscript. 

Literature Cited 

Bergstrom, Jan. 1973. Organization, life, and sys- 
tematics of trilobites. Fossils and Strata 2:1- 

Fritz, W. H. 1 972. Lower Cambrian trilobites from 
the Sekwi Formation type section. Canadian 
Geological Survey Bulletin 212. 

Harrington, H. J. 1959. General description of 
Trilobita, p. 38-1 26 in R. C. Moore (ed.). Trea- 
tise on Invertebrate Paleontology, Part 0, Ar- 
thropoda 1 . Geological Society of America and 
the University of Kansas Press, Lawrence, 

Mount, J. D. 1980. An Early Cambrian fauna 
from the Carrara Formation, Emigrant Pass, 
Nopah Range, Inyo County, California: a pre- 
liminary note, p. 78-80 in Paleontological Tour 
of the Mojave Desert, California-Nevada. 
Southern California Paleontological Society 
Special Publication No. 2. 

Rasetti, Franco. 1948. Lower Cambrian trilobites 
from the conglomerates of Quebec. Journal of 
Paleontology 22:1-24. 

Resser, C. E., and B. F. Howell. 1938. Lower 
Cambrian Olenellus Zone of the Appalachians. 
Geological Society of America Bulletin 49:195- 

Palmer, A. R., and R. B. Halley. 1979. Physical 
stratigraphy and trilobite biostratigraphy of the 
Carrara Formation (Lower and Middle Cam- 
brian) in the southern Great Basin. United 
States Geological Survey Professional Paper 

Walcott, CD. 1910. Olenellus and other genera 
of the Mesonacidae. Smithsonian Miscella- 
neous Collections 53:233-422. 



JUL 91964 



Volume 20 Number 1 pp. 1 51 -1 64 20 June 1 984 

Type specimens of amphibians and reptiles in the 
San Diego Natural History Museum 

Gregory K. Pregill and James E. Berrian 

Department of Herpetology, San Diego Natural History Museum, San Diego, CA 92112 USA 

Publishing a list of type specimens in a collection serves several purposes. For 
those who compile them, the inventory is a propitious means of revealing the taxonomic 
and nomenclatural errors and inconsistencies that routinely creep into a collection. For 
users, the list provides a comprehensive reference to that institution's holdings of 
primary and secondary types, and serves as a guide to the original literature, perhaps 
even yielding insights into the nomenclatural history of the taxa. 

In generating the present account, we made use of the only previous publication 
of type specimens of amphibians and reptiles in the San Diego Natural History Museum, 
compiled by Allan J. Sloan in 1965 (Transactions San Diego Society of Natural History 
(14(1): 1-8). Sloan listed only holotypes, which at the time numbered 48, all of reptiles. 
To make the present list more comprehensive we have included all paratypic material, 
and revised the now expanded list of holotypes. 

The collection currently includes holotypes of 1 frog, 14 lizards and 41 snakes. As 
well, there are several hundred paratypes of 24 additional taxa. Locating all secondary 
types in the collection was a difficult task because many of these specimens had not 
been so designated in the early years of cataloging, nor were they segregated from the 
main collection. Subspecies comprise the majority of primary and secondary types, 
many of which have been synonymized since their original description. Readers familiar 
with the collections of amphibians and reptiles at the San Diego Natural History 
Museum are aware that the geographic emphasis favors the Southwest, Baja California 
and the Gulf of California Islands; this is reflected in the type localities that follow. 

For many years the collections of amphibians and reptiles at the San Diego Natural 
History Museum were closely tied to the activities of Laurence M. Klauber, although 
the full history extends beyond the KJauber era back to the Society's inception in 1 874. 
Klauber began his herpetological studies as an avocation in 1920, conducting most of 
his research and maintaining the collections in the basement of his home. By 1960 he 
had amassed 35 000 specimens. These, as well as his vast library of over 20 000 books 
and reprints, were donated to the San Diego Society of Natural History, whence they 
were housed in the Natural History Museum in Balboa Park. At the time of Klauber's 
death in 1968, the number of specimens of amphibians and reptiles at the museum 
was over 50 000. 

Over the years the collection has had many contributors, among the earliest being 
the celebrated naturalists of the Southwest, Charles R. Orcutt and Frank Stephens. 
Later, during the middle third of this century, the collection benefited considerably 
from the efforts of C. B. Perkins and Charles E. Shaw. Klauber's legendary interest in 
rattlesnakes alone resulted in an enormous series of Crotalus and a substantial repre- 
sentation of reptiles native to the Far West and northern Mexico. Indeed, during the 
1 10-year history of the Society, the museum has benefited from the interests and travels 
of many individuals. Hence, the collections of amphibians and reptiles are flavored 


with series of taxa from, for example, the Galapagos Islands, the Pacific, and the 

As eariy as 1928, Klauber foresaw that someday his collection and that of the 
museum would merge, and he assigned specimen numbers accordingly so* that they 
would not have to be recatalogued at a future date. Consequently, all specimens orig- 
inally in the LMK series have the same numbers in the herpetological catalogue of the 
San Diego Natural History Museum. The collection now has approximately 65 000 
specimens which may be broken down taxonomically as follows: 

Amphibians (Apoda, Caudata, Anura)— 18 families, 60 genera, 175 species 

Turtles— 9 families, 34 genera, 80 species 

Lizards (including amphisbaenians and Sphenodon)— 1 7 families, 1 30 genera, 

360 species 
Snakes— 9 families, 195 genera, 400 species 

These taxa also encompass nearly 2000 skeletons in roughly the same diversity, and 
several hundred preserved hemipenes representing 70 species of snakes (mostly Cro- 
talus) and lizards. 

About This List 

Each species entry begins with the author, date and publication of the original 
description; the specific page number where the description appears is given paren- 
thetically at the end of that citation. This line is followed by the SDSNH number of 
the holotype, sex, type locality and the name of the collector and date that the specimen 
was caught. In nearly all instances, we have tried to preserve the original wording for 
the type locality. Where potential confusion exists because of orthography or archaic 
and vague place names, we have clarified them as needed, inside brackets [ ] or at the 
bottom of the entry under Remarks. When only the paratype(s) of a taxon is represented 
in the SDSNH collection, the holotype and catalogue number of the holding institution 
is given under Remarks. 

Each taxon is listed alphabetically by genus, species and subspecies precisely as it 
was originally designated, that is, irrespective of present usage. For those taxa that have 
been synonymized or whose nomenclature is currently in doubt, this is so noted under 
Remarks with the name and publication of the first authority or re visor. We have made 
no nomenclatural decisions per se, and a particular taxon is considered valid unless 

Abbreviations of museums and personal collections are as follows: 

AMNH: American Museum of Natural History 

BYU: Brigham Young University 

CAS: California Academy of Sciences 

CAS— SU: California Academy of Sciences— Stanford University collection 

CM: Carnegie Museum 

EHT: Edward H. Taylor collection* 

FMNH: Field Museum of Natural History 

LACM: Los Angeles County Natural History Museum 

MCZ: Museum of Comparative Zoology, Harvard University 

SDSNH: San Diego Society of Natural History 

SU: Stanford University Natural History Museum* 

T-S: Edward H. Taylor and Hobart M. Smith, University of Kansas* 

UIMNH: University of Illinois, Museum of Natural History 

UMMZ: University of Michigan, Museum of Zoology 

USNM: National Museum of Natural History 

* Specimens from these collections have been dispersed to other institutions. 



Bufo exsul Myers (1942) 

Occ. Papers Mus. Zool. Univ. Michigan 460:1-13 (p. 3). 

Paratypes. -SDSNH 29098 and 29099 Deep Springs, Inyo Co., California. 
Remarks. — = Bufo boreas exsul 2iCcord\ng to Stebbins (1962, Amphibians of West- 
em North America. Univ. Calif. Press). Holotype: UMMZ 83357 a very large female. 

Hyla regilla cascadae Jameson, Mackay and Richmond (1966) 
Proc. California Acad. Sci. 4th ser. 33(19):55 1-620 (p. 602). 

Paratypes. -SDS^H 44971-44987 '/2 mi S of Bend, Deschutes Co., Oregon. 
Remarks. — Ho\o\y^Q: CAS 101038 an adult male. 

Hyla regilla deserticola Jameson, Mackey and Richmond (1966) 
Proc. California Acad. Sci. 4th ser. 33(1 9):55 1-620 (p. 582). 

Holotype. — SDSiNH 54176 an adult male; San Borjas, Baja California Norte, Mex- 
ico. By David L. Jameson, November 25, 1961. 

Paratypes. — SY>SN\\ 54166-54175 data same as for Holotype. 

Remarks. — =Hyla regilla hypochondriaca according to Duellman (1970, Univ. 
Kansas Mus. Nat. Hist. Monog. No. 1:487). San Borjas reads San Borja on recent maps. 

Hyla regilla pacifica Jameson, Mackey and Richmond (1966) 
Proc. California Acad. Sci. 4th ser. 33(19):55 1-620 (p. 591). 

Paratypes. -Sr>S>HH 53514-53529 4 mi S of Waldport, Lincoln Co., Oregon. 
Remarks. — HoXoXypt: CAS 101007 an adult male. 

Hyla regilla palouse Jameson, Mackey and Richmond (1966) 
Proc. CaUfomia Acad. Sci. 4th ser. 33(19):55 1-620 (p. 599). 

Paratypes. -SUSHH 44715-44718 6 mi SE of La Grande, Union Co., Oregon. 
Remarks. — YioXoXypQ: CAS 100982 an adult male. 

Hyla regilla sierrae Jameson, Mackey and Richmond (1966) 

Proc. California Acad. Sci. 4th ser. 33(19):55 1-620 (p. 605). 

Paratypes. -^\:>S^H 53835-53841 I'/* mi SSE of Tioga Pass Ranger Station. E 
entrance to Yosemite National Park, Mariposa Co., California. 
Remarks. — HoXoly^Q-. CAS 100991 an adult male. 


Anniella geronimensis Shaw (1940) 

Trans. San Diego Soc. Nat. Hist. 9(24):225-228 (p. 225). 

Holotype. — SUS^H 7543 an adult female; San Geronimo Island, Lower California 
[Norte], Mexico. By Margaret [Mrs. Griffing] Bancroft, March 28, 1932. 
Paratypes. — 'S>r>'S>N\\ 7542 data same as for Holotype. 
Remarks. — =Anniella pulchra according to Hunt (1983, Copeia (l):79-89). 

Anolis rivalis Williams (1984) 
Breviora 478:1-22 (p. 7). 

Paratype. -SDSNH 31 163 "Port Utria," [south of Punta Solano, Choco] Colom- 

Remarks. — Holotype: LACM 42124 an adult male. 

Coleonyx variegatus abbotti Klauber (1945) 

Trans. San Diego Soc. Nat. Hist. 10(1 1): 133-2 16 (p. 154). 

Holotype. — SYySnW 34790 an adult male; Proctor Valley, San Diego Co.. Cali- 
fornia. By William Moore, February 28, 1942. 

Paratypes.- As follows: California: Los Angeles Co.: SDSNH 201 1 San Francis- 
quito Plant 2. Riverside Co.: SDSNH 2725 Moreno. San Diego Co.: SDSNH 30 Cot- 


tonwood; SDSNH 843 Foster; SDSNH 16702 Rincon; SDSNH 16988, 32817, 34786 
Mission Gorge; SDSNH 1 6989 Jamui; SDSNH 1 70 1 2 De Luz (says "Sentenac Canyon" 
in catalogue); SDSNH 21249 El Capitan; SDSNH 24050 San Pasqual; SDSNH 25303, 
34666 Jacumba; SDSNH 27770 Foot Agua Tibia Mt.; SDSNH 32797 Pala; SDSNH 
32821-32822 Black Mtn. near La Mesa. Baja California Norte, Mexico: SDSNH 2593 
Ensenada; SDSNH 5265-5266, 15970-15971, 27726, 30295 Cedros Island; SDSNH 
6553 65 mi SE of Tecate; SDSNH 24390 San Jose (lat. 32°). 

Coleonyx variegatus bogerti Klauber (1943) 

Trans. San Diego Soc. Nat. Hist. 10(1 1):133-216 (p. 176). 

Ho/otvpe. — SDSNH 32486 an adult male; Xavier, Pima Co., Arizona. By Lee W. 
Arnold, July 17, 1939. 

Paratypes.— None designated. 

Coleonyx variegatus utahensis Klauber (1943) 

Trans. San Diego Soc. Nat. Hist. 10(1 1):133-216 (p. 171). 

Holotype. — SDSNH 35792 an adult male; Watercress Spring, Washington Co., 
Utah. By Dr. Ross Hardy, April 16, 1941. 

Paratypes. — SDSNH 36021-36024 data same as for Holotype. 

Phyllodactylus angelensis Dixon (1966) 

Proc. Cahfomia Acad. Sci. 4th ser. 33(13):4 15-452 (p. 444). 

Holotype. — SDSNH 19996 an adult male; north end of Isla Angel de la Guarda, 
Baja California [Norte], Mexico. By Allan J. Sloan, March 22, 1963. 

Paratype. — SDSNH 50851 Isla Pond, [Baja California Norte, Mexico]. 

Phyllodactylus apricus Dixon (1966) 

Proc. California Acad. Sci. 4th ser. 33(1 3):4 15-452 (p. 450). 

Holotype. — SDSNH 44623 an adult male; Isla Las Animas, Baja California [Sur, 
Mexico]. By Chris Parrish and G. E. Lindsay, June 27, 1964. 

Paratypes.-SDSNH 44620-44622, 44624, 50830-50842, 50844-50849 data same 
as for Holotype. 

Phyllodactylus bugastrolepis Dixon (1966) 

Proc. Cahfomia Acad. Sci. 4th ser. 33(13):4 15-452 (p. 447). 

Paratypes.-SDSNH 44604-44607, 44611-44613, 50792, 50793 Isla Catalina, 
Baja California [Sur, Mexico]. 

Remarks. — HoXoXyTpe: CAS 98485 an adult female. 

Phyllodactylus homolepidurus nolascoensis Dixon (1964) 
New Mexico State Univ. Sci. BuH. 64(1): 1-1 39 (p. 42). 

Paratypes.-SDSNH 6840 and 6841 Isla San Pedro Nolasco, Sonora, Mexico. 
Remarks. — Ho\o\ype: CAS 50552 an adult male. 

Phyllodactylus partitus Dixon (1966) 

Proc. California Acad. Sci. 4th ser. 33(13):4 15-452 (p. 445). 

Paratypes.-SDSNH 6834-6836, 39258, 39649, 40508, 50820-50822 Isla Partida 
(N), Baja California [Norte, Mexico]. 

Remarks. — HoXoXype: CAS 98429 an adult male. 

Phyllodactylus santacruzensis Dixon (1966) 

Proc. California Acad. Sci. 4th ser. 33(13):4 15-452 (p. 449). 

Paratypes.-SDSNH 50872, 50873 Isla Santa Cruz, Baja California [Sur, Mexico]. 
Remarks. -Holotype: CAS 98468 an adult female. 

Phyllodactylus xanti acorius Dixon (1966) 

Proc. Cahfomia Acad. Sci. 4th ser. 33(13):4 15-452 (p. 442). 

Paratypes.-SDSNH 50827-50829 Isla San Diego, Baja California [Sur, Mexico]. 


Remarks. — =Phyllodactyhis nocticolus acohiis according to Murphy (1983, Occ. 
Papers California Acad. Sci. 137:1-48). Holotype: CAS 98451 an adult male. 

Phyllodactylus xanti angulus Dixon (1966) 

Proc. California Acad. Sci. 4th ser. 33(13):4 15-452 (p. 433). 

Paratypes.-SX:>^NH AAbll , 44678, 50868, 50869 Isla Salsipuedes, Baja California 
[Norte, Mexico]; SDSNH 44238, 44239, 50833, 50856-50859 [Isla] San Lorenzo, [Baja 
California Norte, Mexico]. 

Remarks. — =PhyUodactylus nocticolus angulus according to Murphy (1983, Occ. 
Papers California Acad. Sci.' 137:1-48). Holotype: CAS 98477 an adult male. 

Phyllodactylus xanti circus Dixon (1966) 

Proc. California Acad. Sci. 4th ser. 33(13):4 15-452 (p. 439). 

Holotype. — SUS^H 50814 an adult female; Isla Ildefonso [Baja California Sur, 
Mexico]. By Charles E. Shaw, April 2, 1962. 

Paratypes.-^X:>Sn\\ 50809-50812, 50815-50817 data same as for Holotype. 

Remarks. — ^Phyllodactylus nocticolus circus according to Murphy (1983, Occ. 
Papers California Acad. Sci. 137:1-48). 

Phyllodactylus xanti estebanensis Dixon (1966) 

Proc. California Acad. Sci. 4th ser. 33(13):4 15-452 (p. 437). 

Paratypes.-SX:>S^H 50852 and 50853 Isla Tiburon [Gulf of California, Mexico]; 
50870 and 50871 Isla San Esteban [Gulf of California, Mexico]. 

Remarks. — = Phyllodactylus nocticolus estebanensis according to Murphy (1983, 
Occ. Papers California Acad. Sci. 137:1-48). Holotype: CAS 98481 an adult male. Isla 
Tiburon and Isla Esteban belong to the state of Sonora, 

Phyllodactylus xanti nocticolus Dixon (1964) 

New Mexico State Univ. Sci. Bull. 64(1): 1-1 39 (p. 55). 

Holotype. SY^S^W 34824 an adult male; Agua Caliente Hot Springs, San Diego 
Co., California. By Laurence M. Klauber [Charles Shaw and Paul Breese]. Preserved 
March 8, 1942. 

Paratypes.-¥rom San Diego County, as follows: SDSNH 2952, 18633, 32955, 
35476-35490, 40073 Yaqui Well; SDSNH 18632, 33247 Sentenac Canyon; SDSNH 
1 892 1 Yaqui Pass; SDSNH 33672-33674 Palm Canyon; SDSNH 34825-34827, 43786, 
43787 Topotypes; SDSNH 38034, 38041 4 mi W of Vallecitos Stage Station. 

Remarks. — = Phyllodactylus nocticolus nocticolus according to Murphy (1983, Occ. 
Papers California Acad. Sci. 137:1-48). 

Phyllodactylus xanti sloani Bostic (1972) 

Trans. San Diego Soc. Nat. Hist. 16(10):237-263 (p. 252). 

Holotype. -SDSNH 45895 an adult female; 23.5 mi SE of El Rosario, Baja Cal- 
ifornia Norte, Mexico. By Dennis L. Bostic, March 31, 1969. 

Paratypes.- AW from Baja California Norte as follows: SDSNH 45896 1.8 mi NW 
of Puerto de San Carlos: SDSNH 45897 and 45898 5.4 mi W of Punta Canoas; SDSNH 
45899 and 45900 Arroyo San Jose; SDSNH 4590 1-45907 Las Palomas; SDSNH 45908 
EI Cardon; SDSNH 45909-45912 5 mi N of San Javier. 

Remarks. — = Phyllodactylus nocticolus sloani according to Murphy (1983, Occ. 
Papers California Acad. Sci.' 137:1-48). SDSNH 45912 was "sent to Mexico, 1971" 
[presumably to Fauna Silvestre, Mexico City]. 

Sauromalus australis Shaw (1945) 

Trans. San Diego Soc. Nat. Hist. 10(15):269-306 (p. 286). 

Holotype. — SDSNH 30170 an adult male; San Francisquito Bay, Baja California, 
Mexico. By Robert S. Hoard, July 30, 1938. 

Paratypes. — W\ are from Baja California Sur as follows: SDSNH 1 7707 Comondu; 
SDSNH 17708 La Paz; SDSNH 30168 Loreto; SDSNH 30169 33 mi N of Canipole. 


Sauromalus klauberi Shaw (1941) 

Trans. San Diego Soc. Nat. Hist. 9(28):285-288 (p. 285). 

//o/ory;?£'. — SDSNH 6859 an adult male; Santa Catalina Island, Gulf of California 
[Baja California Sur], Mexico. By J. R. Pemberton, December 14, 1931. * 

Paratypes. — SDSHW 6860 and 6861 data same as for Holotype. 

Remarks. — ^Sauromulus ater klauberi according to Avery and Tanner (1964, 
BYU Science Bull. 5(3): 1). 

Sauromalus obesus tumidus Shaw (1945) 

Trans. San Diego Soc. Nat. Hist. 10(15):269-306 (p. 292). 

//o/or.vp^— SDSNH 27323 an adult male; Telegraph Pass, Gila Mountains, Yuma 
Co., Arizona. By Laurence M. Klauber, June 15, 1937. 

Paratypes.-ST:>S^H 8613, 27551, 33170-33175, 33224, 33225, 34141, 35090 
data same as for Holotype (Gila Mountains, Arizona). 

Uma notata cowlesi Heifetz (1941). 
Copeia 1941(2):99-1 1 1 (p. 104). 

Paratypes. — 'SDS^W 16460-16464 Punta Penasco, Sonora, Mexico. 
Remarks. — = Uma notata rufopimctata Cope according to Norris (1958, Bull. Amer. 
Mus. Nat. Hist. 1 14(3):25 1-326). Holotype: CAS 53370 an adult male. 

Uta stansburiana klauberi Lowe and Norris (1955) 
Hereptologica ll(2):89-96 (p. 91). 

Holotype. — SDS^H 6642 an adult male; San Esteban Island, Gulf of CaUfomia, 
Sonora, Mexico. By J. R. Pemberton, January 1 1, 1932. 

Paratypes.-Sr>%Nn 3968-3971, 6640, 6641 San Esteban Island, Sonora, Mexico. 

Remarks. — = Uta stansburiana tavlori Smith according to Ballinger and Tinkel 
(1972, Misc. Pub. Mus. Zool. Univ. Michigan (145): 1-83). 

Xantusia arizonae Klauber (1931) 

Trans. San Diego Soc. Nat. Hist. 7(1): 1-1 6 (p. 3). 

Holotype. SY^SNYi 5433 an adult female; 1 mi S of Yamell, Yavapai Co., Arizona. 
By Philip M. Klauber and Laurence M. Klauber, August 21, 1931. 

Paratypes.-S\:>^^H 5434-5438 data same as for Holotype; SDSNH 5451 and 
5452 bom (dead) of Holotype in captivity; SDSNH 5450 unborn embryo from SDSNH 
5436: SDSNH 5453 unborn embryo from SDSNH 5434. 

Remarks. — =Xantusia vigilis arizonae according to Bezy (1967 Copeia (3):653- 
661). SDSNH 5434 was sent to J. R. Slevin at the California Academy of Sciences 
December 14, 1931. 


Arizona elegans blanchardi Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 10(17):31 1-398 (p. 328). 

Paratype. — SDSNH 35343 Schramm, Yuma Co., Colorado. 

Remarks. -WoXoXy^Q: SU 10393 a young adult male; now CAS-SU 10393. 

Arizona elegans Candida Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 10(17):31 1-398 (p. 364). 

Holotype. — SDSNH 34191 an adult male; Kramer Hills, 6 mi S of Kramer Junction 
on US 395, San Bernardino Co., California. By James Deuel, June 16, 1941. 

Paratvpes. — M\ are from near the type locality in San Bernardino Co., California, 
as follows: Adelanto: SDSNH 2725 1 , 34 1 87, 34 1 90 1 6 mi N of; SDSNH 28848, 33980 
8 mi N of; SDSNH 3 1 9 1 7 1 2 mi S of; SDSNH 3 1 94 1 , 3 1 942, 35 109 3 mi N of; SDSNH 
33323 6 mi SW of; SDSNH 338 1 2 6 mi N of; SDSNH 3398 1 , 35093 4 mi S of; SDSNH 
33982 14 mi N of; SDSNH 34107 13 mi N of; SDSNH 34150, 35594 2 mi N of 
Kramer Hills: SDSNH 28846, 31700, 33795, 35149. Kramer Junction: SDSNH 31766 


5 mi N of; SDSNH 31940, 33832, 33983, 34019 8 mi S of; SDSNH 31959 20 mi S 
of; SDSNH 33826, 34189 3 mi S of; SDSNH 33888, 34164, 34192 5 mi S of; SDSNH 
33975, 34017, 35536 7 mi S of; SDSNH 33977, 33978 10 mi S of; SDSNH 33979 12 
mi S of; SDSNH 34165 6 mi of; SDSNH 34184 1 mi W of; SDSNH 34185 1 mi E of; 
SDSNH 34186 1 1 mi S of; SDSNH 35151, 35537 9 mi S of SDSNH 35654 6.5 mi 

Arizona elegans eburnata Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 10(17):31 1-393 (p. 350). 

Holotype. — ^Y^SNH 33094 a young male; Benson's Dry Lake, San Diego Co., 
California. By James Deuel, June 5, 1940 [preserved]. 

Paratvpes. — M\ are from near the type locality in San Diego Co., California, as 
follows: SDSNH 4454, 23854, 23914, 23915, 25437, 25438, 26914, 26939-26942, 
27308, 27383, 27405, 29300, 33094, 33095, 17026 Topotypes; SDSNH 4862 Beatty 
Ranch, Borrego Valley; SDSNH 5136, 21 108, 21 121 Borrego Valley; SDSNH 23024, 
23774-23776, 23852, 23853, 26814, 29301, 29487, 32035 The Narrows; SDSNH 
23773 5 mi E of The Narrows; SDSNH 26732 2 mi S of Borrego Post Office; SDSNH 
27331 3 mi W of Benson's Dry Lake. 

Remarks. -SiDS^H 4454, 17026, 21 108 were exchanged with H. M. Smith, Jan- 
uary 14, 1957 and are presumed to be at the University of Illinois Museum of Natural 
History. KJauber also gave as paratypes SDSNH 26056 and 26057, but these are actually 
specimens of Pituophis melanoleucus deserticola and Crotalus viridis oreganus, respec- 
tively; the source of this error is unknown. 

Arizona elegans noctivaga Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 10(17):31 1-398 (p. 343). 

Holotype. — SDSNH 34188 a young adult male; 8 mi N of Owlshead, Pinal Co., 
Arizona. By Charles E. Shaw and Laurence M. Klauber, May 31, 1941. 

Paratypes.— AW are from Arizona, as follows: Gila Co.: SDSNH 34438 7 mi SE 
of Globe. Maricopa Co.: SDSNH 15835, 23925, 23926 Sentinel; SDSNH 25829, 260 1 
Mesa; SDSNH 26913 5 mi S of Wickenburg; SDSNH 26943 Cactus Gardens; SDSNH 
32781 Gila Bend; SDSNH 34331 Stanwix. Pima Co.: SDSNH 13724, 29222 Tucson: 
SDSNH 1 7949 1 1 mi N of Tucson; SDSNH 1 7950, 17951 4V2 mi N of Tucson; SDSNH 
17952 2 mi N of Tucson; SDSNH 27177 13 mi N of Tucson; SDSNH 27178 4 mi 
N of Sahuarita; SDSNH 32293 Martinez Hill; SDSNH 32521 1 1 mi S of Tucson; 
SDSNH 32714 2 mi NE of Tanque Verde Ranch; SDSNH 34018 14 mi N of Tucson. 
Pinal Co.: SDSNH 21492, 21493 Picacho; SDSNH 27180 Rorence; SDSNH 32323 8 
mi W of Casa Grande; SDSNH 34104 3 mi W of Superior; SDSNH 34332 Oracle 
Junction. Yavapai Co.: SDSNH 1 7623 1 mi S of Congress Junction. Yuma Co.: SDSNH 
34526 5 mi E of Salome. 

Arizona elegans pacata Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 10(1 7):31 1-398 (p. 379). 

Holotype. -SDSNH 17652 an adult male; Santo Domingo (lat. 25°30'N), Baja 
California Sur, Mexico. By Frank F. Gander, November 16, 1941. 

Arizona elegans philipi Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 10(1 7):31 1-398 (p. 333). 

Holotype. — SDSNH 34456 an adolescent male; 10 mi E of Winslow, Navajo Co., 
Arizona. By Charles E. Shaw and Carl Engler, July 29, 1941. 

Paratypes. -SDSNH 20990 Two Guns, Coconino Co., Arizona; SDSNH 34426 
data same as for Holotype. 

Charina bottae umbratica Klauber (1943) 

Trans. San Diego Soc. Nat. Hist. 10(7):83-90 (p. 83). 

Holotype. — SDSNH 12101 an immature male; Fern Valley, near Idyllwild, Riv- 
erside Co., California. By Clyde Searl, July 1, 1929. 


Remarks.— The validity of C. b. umbratica as a subspecies has been questioned 
on several occasions, whereas in other instances full species status has been proposed. 
The taxonomic history is treated by Stewart (1977, SSAR Cat. Amer. Amph. Rept. 

Chilomeniscus stramineus esterensis Hoard (1939) 

Pomona College Jour. Ent. and Zool. 31(4):45-46 (p. 45). 

//o/o/V/?^— SDSNH 30368 an adult male; Estero[s] Salina, Lower [Baja] California, 
Mexico [24°36'N, 1 1 1°49'W]. By R. S. Hoard, July 10, 1938. 

/'ararv/?6'5.-SDSNH 30364-30367, 30369, 30370 data same as for Holotype. 

Chionactis occipitalis talpina Klauber (1951) 

Trans. San Diego Soc. Nat. Hist. 1 1(9): 141-204 (p. 172). 

Paratypes.-SY:>S^\l 39520, 39521 10 mi N of Goldfield, Esmerelda Co., Nevada. 
Remarks. — WoXoXype: CAS 81364 an adult male. 

Chionactis palarostris organica Klauber (1951) 

Trans. San Diego Soc. Nat. Hist. 1 1(9): 141-204 (p. 178). 

Holotype. —SD^^W 40673 an adult male; on the Sonoyta-Ajo road, 9 mi N of the 
U.S. -Mexican border, in Organ Pipe Cactus National Monument, Pima Co., Arizona. 
By William R. Supernaugh and Grover E. Steele, May 22, 1950. 

Crotalus cerastes cercobombus Savage and Cliff (1953) 
Nat. Hist. Misc. 1 19:1-7 (p. 2). 

Paratvpes.—M\ from Arizona, as follows: Maricopa Co.: SDSNH 979-981 18 mi 
W of Phoenix; SDSNH 17072 10 mi E of Gila Bend; SDSNH 22410-22412 Desert 
Wells; SDSNH 23879, 23888-23890, 25553-25555, 25854-25857 near Mesa; SDSNH 
26915 1 mi S of Morristown; SDSNH 39082 26 mi N of Ajo; SDSNH 39088 1 mi W 
of Tartron [sic] (see Remarks); SDSNH 40893 vicinity of Phoenix. Pima Co.: SDSNH 
2324, 2325 Sells; SDSNH 38660 Sonoita (see Remarks). Pinal Co.: SDSNH 17068- 
17071, 17073 3 mi SE of Picacho; SDSNH 25499 5 mi W of Casa Grande; SDSNH 
41110-41116,41 362, 4 1 363 4 mi E of Coolidge; SDSNH 4 1 24 1 4 mi S of Coolidge. 

Remarks. — WoXoXyi^e: SU 7287 an adult male; now CAS-SU 7287. The correct 
spelling is Tarton. Sonoita is in Santa Cruz Co. 

Crotalus cerastes laterorepens Klauber ( 1 944) 

Trans. San Diego Soc. Nat. Hist. 10(8):9 1-126 (p. 94). 

Holotype. — SDSNH 34074 an adult male; The Narrows, San Diego Co., California. 
By Cyrus B. Perkins and Charles Shaw, June 6, 1941. 

Par at vpes.— All are from within a 13-mile radius of the type locality as follows: 
SDSNH 1762, 1858, 2209, 2210, 4571, 4572, 4645-4647, 4827, 4875, 4931, 4958, 
5052-5055, 5173-5175, 9507, 21098, 21426, 22275, 22357, 23009, 23233, 23640, 
23858, 23860, 23917, 23954-23956, 23998, 24006, 24019, 24020, 25423, 25445, 
26729-26731, 26823, 26847, 26848, 26857, 26865, 26866, 26937, 26938, 27240, 
28113, 28228, 28682, 28683, 28728, 28750, 29085, 29118, 29119, 29271, 29277, 
29898, 30719, 31930, 31999, 32307, 32977, 32978, 33044, 33058, 33059, 33123, 
33333-33335, 33342, 34035, 34176-34179, 34351, 34570, 35179, 35187, 35188, 35305, 
35557-35559, 35597, 35634, 35635. 

Crotalus confluentus abyssus Klauber (1930) 

Trans. San Diego Soc. Nat. Hist. 6(3):95-144 (p. 1 14). 

Holotype. — SDSNH 2216 an adult male; Tanner Trail, 300 feet below south rim 
of the Grand Canyon, Coconino Co., Airzona. By E. D. McKee, September 15, 1929. 

Paratypes.— None designated. 

Remarks. —=Crotalus viridis abvssus according to Klauber (1936, Trans. San Diego 
Soc. Nat. Hist. 8(20): 185-276). 


Crotalus confluentus kellyi Amaral (1929) 

Bull. Antivenin Inst. Amer. 2(4):86-97 (p. 91). 

Holotvpe. — SX^S^H 194 a male; Needles [San Bernardino Co.], California. By Mr. 
O. R. West, July 11, 1926. 

Paratype. — SV>S,N\\ 195 data same as for Holotype. 

Remarks. — =Cwtalus s. scutulatus according to Klauber (1930. Trans. San Diego 
Soc. Nat. Hist. 6(3):95-144. 

Crotalus confluentus lutosus Klauber (1930) 

Trans. San Diego Soc. Nat. Hist. 6(3):95-144 (p. 100). 

Holotvpe. — SDSNH 1814a young adult male; 10 mi NW of Abraham on the road 
to Joy, Millard Co., Utah. By Cyrus B. Perkins, May 12, 1929. 

Paratvpes.-SDSnYl 1800-1813, 1815, 1816 20 mi NW of Delta, Millard Co., 

Remarks. — =Crotalus viridis lutosus according to Klauber (1936, Trans. San Diego 
Soc. Nat. Hist. 8(20): 195-276). 

Crotalus confluentus nuntius Klauber (1935) 

Trans. San Diego Soc. Nat. Hist. 8(13):75-90 (p. 78). 

//c»/o/y/7£'. — SDSNH 3105 an adult male; Canyon Diablo, Coconino Co., Arizona. 
By R. L. Borden, August 9, 1930. 

Remarks. —=CrotaIus viridis nuntius according to Klauber (1936, Trans. San Diego 
Soc. Nat. Hist. 8(20): 185-276). 

Crotalus durissus culminatus Klauber (1952). 
Bull. Zool. Soc. San Diego 26:1-143 (p. 67). 

Para/yp^.— SDSNH 43403 Hacienda El Sabino, near Uruapan, Michoacan, Mex- 

Remarks. — YioXoXyTpQ-. FMNH 126616 (formerly EHT 5224) a juvenile female. 
SDSNH 43403 was formerly EHT 5233. 

Crotalus mitchelli angelensis Klauber (1963) 

Trans. San Diego Soc. Nat. Hist. 13(5):73-80 (p. 75). 

//o/oO'P^.- SDSNH 51994 an adult male; 4 mi SE of Refugio Bay, Isla Angel de 
la Guarda, Gulf of California, Mexico. By Dr. Reid Moran, March 22, 1963. 

Paratypes.-^U^NH 19717, 19718, 19990-19995, 44358, 51991-51993, 51995, 
51996. All are from Isla Angel de la Guarda. 
Remarks. — Ende:rx\\c to Isla Angel de la Guarda. 

Crotalus mitchellii [sic] muertensis Klauber (1949) 

Trans. San Diego Soc. Nat. Hist. 1 1(6):61-1 16 (p. 97). 

//o/c»0'P^— SDSNH 37447 an adult male; El Muerto Island, Gulf of California, 
Mexico. By Charles H. Lowe, Jr., June 6 or 7, 1946. 

Paratypes.-SDS^H 37442-37444, 37446, 37448, 37449, 38040 El Muerto Island. 

Remarks. — E[ Muerto Island = Isla Miramar in the San Luis group, Baja California 
Norte. The correct spelling is mitchelli. 

Crotalus molossus estebanensis Klauber ( 1 949) 

Trans. San Diego Soc. Nat. Hist. 1 1(6):61-1 16 (p. 104). 

//o/o/V'/?^.— SDSNH 26792 an adult female; San Esteban Island, Gulf of California, 
Mexico. By an expedition under Capt. G. Allan Hancock, April 17. 1937 (preserved). 
Remarks. — Endemic to Isla San Esteban. 

Crotalus ruber lorenzoensis Radcliffe and Maslin (1975) 
Copeia 1975(3):490-493 (p. 490). 

//o/o/v/?^— SDSNH 46009 an adult male; San Lorenzo Sur Island, Gulf of Cali- 
fornia, Baja California Norte, Mexico. By Charles E. Shaw, May 23, 1966. 


Paratypes. -SUSNH 6605, 45052, 45053, all from San Lorenzo Island Sur. 
Remarks. — Endemic to San Lorenzo. 

Crotalus triseriatus anahuacas Gloyd ( 1 940) 

Chicago Acad. Sci. Special Publ. No. 4:1-270 (p. 91). * 

Paratvpes. — ^DSHYi 43404 43 km N of Tres Cumbres, Morelos, Mexico. 
Remarks. -HoXoXyipQ: MCZ 33681 a female. SDSNH 43404 was formerly T-S 

Crotalus triseriatus aquilus Klauber (1952) 

Bull. Zool. Soc. San Diego 26:1-143 (p. 24). 

Paratypes.-SUS^H 3496-3501, 6575-6577 vicinity of Alvarez, San Luis Potosi, 
Mexico (Topotypes). 

Remarks. — =^Crotallus aquilus according to Harris and Simmons (1978, Bull. 
Maryland Herp. Soc. 14(3):105-21 1). Holotype: MCZ 27843 an adult female. 

Crotalus vegrandis Klauber (1941) 

Trans. San Diego Soc. Nat. Hist. 9(30):333-335 (p. 334). 

Paratypes.SDS^H 34607 Maturin Savannah, near Uracoa Monagas, Venezuela. 
Remarks. -HoXoXype: CM 17384 an adult male. SDSNH 34607 was formerly CM 


Crotalus viridis caliginis Klauber (1949) 

Trans. San Diego Soc. Nat. Hist. 1 1(6):61-1 16 (p. 90). 

//o/o/.vp^. — SDSNH 2800 an adult male; South Coronado Island off the northwest 
coast of Baja California, Mexico. By E. H. Quayle, June 2, 1930. 

Paratypes.-SDS^U 2801-2804, 4924-4926, 7538-7540, 1 1 177, 1 1 178, 1371 1- 
13715, 20077, 20078 "All probably came from South Coronado Island, Baja California 
Norte, Mexico." 

Remarks. — ¥jno^n only from the type locality. 

Crotalus willardi meridionalis Klauber (1949) 

Trans. San Diego Soc. Nat. Hist. 1 1(8): 12 1-1 40 (p. 131). 

Holotype. — SDSNW 6569 an adult female; Coyotes ["on the railroad to El Sato"], 
elevation 8000', Durango, Mexico. By Edmund Heller and Charles M. Barber, August 

Remarks. — S>r>SHY{ 6569 was one of two specimens obtained from the Field Mu- 
seum, both of which were numbered FMNH 1493. 

Hypsiglena nuchalatus W. Tanner (1943) 
Great Basin Nat. 4(1 & 2):49-54 (p. 49). 

Paratypes.-ST:>SNH 20233, 20293, 22501 Visalia, Tulare Co., California. 

Remarks. — =Hvpsiglena torquata nuchulata according to Bogert and Oliver ( 1 945, 
Bull. Amer. Mus. Nat. Hist. 83(6):297-426 (p. 381)). Holotype: BYU 3008 a small 

Hypsiglena ochrorhynchus [sic] klauberi W. Tanner (1944) 
Great Basin Nat. 5(3 & 4):25-92 (p. 71). 

//o/ory/7^. — SDSNH 20228 a male; South Cornoado [=Coronado] Island, Lower 
California' [Baja California Norte]. By Philip M. Klauber, June 11, 1933. 

Remarks. — =Hypsiglena torquata klauberi according to Bogert and Oliver (1945, 
Bull. Amer. Mus. Nat. Hist. 83(6):297-426). A review of the nomenclatural problem 
surrounding ochrorhvncha Cope vs. torquata Giinther is given by Hardy and Mc- 
Diarmid (1969:169, Univ. Kansas Publ. Mus. Nat. Hist. 18(3):39-252). 

Hypsiglena ochrorhynchus [sic] tortugaensis W. Tanner (1944) 
Great Basin Nat. 5(3-4):25-92 (p. 69). 

Paratypes.—^DS^H 4074 Tortuga Island [Baja California Sur, Mexico]. 


Remarks. — =Hypsiglena torquata tortugaensis according to Bogert and Oliver ( 1 945, 
Bull. Amer. Mus. Nat. Hist. 83(6):297-426). Holotype: CAS 51460 a female (by in- 
ference). See also Remarks under H. o. klauberi. 

Hypsiglena torquata catalinae W. Tanner (1966) 

Trans. San Diego Soc. Nat. Hist. 14(1 5): 189-1 96 (p. 192). 

//(9/o/\'/7£'. — SDSNH 44680 an adult male; Santa Catalina Island, approximately 
25°38'N, ilO°47'W, Gulf of California, Baja California [Sur], Mexico. By George E. 
Lindsay, June 25, 1964. 

Paratypes.-SU^^H 44376, 44681 Topotypes. 

Lampropeltus zonata pulchra Zweifel (1952) 
Copeia 1952(3): 152-1 68 (p. 162). 

Holotype. — S,r>S>N\\ 38667 an adult male; near Crater Camp [450 ft], Santa Monica 
Mountains, Los Angeles Co., California. 

Remarks. — Date and collector unknown. Apparently received from the San Diego 
Zoo, preserved August 2, 1947. 

Leptotyphlops humilis cahuilae Klauber (1931) 

Trans. San Diego Soc. Nat. Hist. 6(23):333-352 (p. 339). 

//o/o/yp6'. — SDSNH 2637 an adult; Yaqui Well by the County Road Camp, San 
Diego Co., California. By Laurence M. Klauber, May 15, 1930. 
Paratypes.—^onQ designated. 
Remarks. — Sex of Holotype not given. 

Leptotyphlops humilis lindsayi Murphy (1975) 

Proc. California Acad. Sci. ser. 4, 40(5):93-107 (p. 96). 

Holotype.— SUSNH 44386 an adult female; Marquer Bay, Isla Carmen, Baja 
California (Sur), Mexico. By Charles E. Shaw and George E. Lindsay, April 4, 1962. 
Remarks. — Yjivo^jn only from Holotype. 

Lichanura roseofusca gracia Klauber (1931) 

Trans. San Diego Soc. Nat. Hist. 6(20):305-318 (p. 307). 

Holotype. — SDSNH 2995 a young female; Randsburg, Kern Co., California. By 
Lucile Rector, June, 1930. 

Remarks. — =Lichamira trivirgata gracia according to Miller and Stebbins (1964: 
189 The Lives of Desert Animals in Joshua Tree Nat'l Monument, Univ. Calif. Press 
vi + 452 p.) and others since, all without comment. For a discussion see Yingling 
(1982, SSAR Cat. Amer. Amph. Rept. 294.1). 

Masticophis bilineatus lineolatus Hensley (1950) 

Trans. Kansas Acad. Sci. 53(2):270-288 (p. 272). 

//o/oryp<?.-SDSNH 43402 an adult male; 12.9 mi S and 5 mi E of Ajo, Pima Co., 

Remarks. -SDS^H 43402 was formerly UIMNH 561 1. 

Masticophis bilineatus slevini Lowe and Norris (1955) 
Herpetological 1 1(2):89-96 (p. 93). 

Holotype. — SDSNH 3826 an adult female; San Esteban Island, Gulf of California, 
Sonora, Mexico. By Mrs. Griffing [Margaret] Bancroft, April 18. 1930. 

Paratype. — SDSNH 41571 San Esteban Island, Gulf of California, Sonora, Mexico. 

Phyllorhynchus browni lucidus Klauber (1940) 

Trans. San Diego Soc. Nat. Hist. 9(20): 1 95-2 14 (p. 202). 

Holotype. — SDSNH 28819 a juvenile female; Enchanto Valley, 7 mi W of Cave 
Creek, Maricopa Co., Arizona. By V. Housholder, May 21, 1938. 

Phyllorhynchus decurtatus nubilus Klauber ( 1 940) 

Trans. San Diego Soc. Nat. Hist. 9(20): 195-2 14 (p. 197). 


Holotype. — SDSNH 32493 an adult male; Xavier (Weisner's Ranch), Pima Co., 
Arizona. By Lee Arnold, July 16, 1939. 

Paratvpes. — From Arizona as follows: Pima Co.: SDSNH 29216 2 mi N of San 
Xavier Mission; SDSNH 29287 4 mi N of San Xavier Mission; SDSNH 3^2273 Ajo 
Junction (Escuela); SDSNH 32289 4 mi S of Ajo Junction; SDSNH 32290 2 mi S of 
Ajo Junction near Tucson; SDSNH 32468 east base of ''A" Mountain; SDSNH 32494 
5 mi N of Tucson. Pinal Co.: SDSNH 32274 26 mi N of Tucson. 

Remarks. — ¥Aauber noted in his catalogue that SDSNH 29287 and 32468 were 
"traded to Slevin, 1941"; these are now CAS specimens. 

Phyllorhynchus decurtatus perkinsi Klauber (1935) 
Bull. Zool. Soc. San Diego 12:1-31 (p. 11). 

//o/oO'P^. — SDSNH 23757 an adult male; [Benson's] Dry Lake 3 mi W of Imperial 
Co. Line on Narrows, Kane Springs Road, San Diego Co., California. By Laurence M. 
Klauber, May 4, 1935. 

Paratvpes— ^D^^U 22260, 22261, 22295, 22297, 22298, 22741, 22954, 22955, 
23027, 23386, 23387, 23750-23756, 23758-23761, 23815, 23846-23848, 2391 1-23913, 
23918-23923, 23951, 23995-23997 all from very near the type locality. 

Pituophis catenifer bimaris Klauber ( 1 946) 

Trans. San Diego Soc. Nat. Hist. 1 1(1): 1-40 (p. 7). 

//o/on^/?^'. — SDSNH 3262 1 an adult male; Santa Gertrudis, near El Arco (lat. 28°N), 
Baja California, Mexico. By Robert S. Hoard, August, 1939. 

Paratvpes.- A\\ from Baja California, Mexico, as follows: SDSNH 2934, 2935 El 
Refugio (northeast of Magdalena Bay); SDSNH 32523 18 mi N of Punta Prieta (lat. 
27°N); SDSNH 3813, 11553 San Ignacio; SDSNH 31032 Bahia Thurtoe (1 mi S of 
Bahia Tortuga); SDSNH 17562 and 17563 Rancho Las Flores (12 mi E of El Arco); 
SDSNH 1129, 1181 El Marmol. 

Remarks. — =Pituophis melanoleucus bimaris according to Smith and Kennedy 
(1951, Hereptologica 7(3):93-96). 

Pituophis catenifer coronalis Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 1 1(1): 1-40 (p. 19). 

Holotype.-SUSNW 20229 an adult female; South Coronado Island, Baja Cali- 
fornia Norte, Mexico. By Philip M. Klauber, June 11, 1933. 

Paratype. — SD^^Yi 1 1365 Coronado Islands, Baja California Norte, Mexico (re- 
stricted to South Coronado Island by Klauber, ibid p. 20). 

Remarks. — =Pituophis melanoleucus coronalis according to Smith and Kennedy 
(1951, Herpetologica 7(3):93-96). 

Pituophis catenifer fuliginatus Klauber (1946) 

Trans. San Diego Soc. Nat. Hist. 11(1): 1-40 (p. 14). 

//o/o/vp£'. — SDSNH 17449 a young adult female; San Martin Island, off the west 
coast of Baja California, Mexico. By Lewis W. Walker, July 1 1, 1939. 

Paratypes.-^DS^H 17463, 17464 Topotypes. 

Remarks. — =Pituophis melanoleucus fuliginatus according to Smith and Kennedy 
(1951, Herpetologica 7(3):93-96). 

Pituophis catenifer pumilus Klauber ( 1 946) 

Trans. Soc. Diego Soc. Nat. Hist. ll(2):41-48 (p. 41). 

Holotype. — SDS^H 17238 a young adult male; Santa Cruz Island, Santa Barbara 
Co., California. By Norman Bilderback, May 5, 1938. 

Remarks. — =Pituophis melanoleucus pumilis according to Smith and Kennedy 
(1951, Herpetologica 7(3):93-96). 

Rhinochelus lecontei clarus Klauber (1941) 

Trans. San Diego Soc. Nat. Hist. 9(29):289-332 (p. 308). 


Holotvpe. — SDSNH 3 1440 an adult male; Borrego Valley, 2 mi N of The Narrows, 
San Diego Co., California. By Richard Neil, May 7, 1939. 

Paratvpes.- As follows: San Diego County: SDSNH 2631, 1 1288, 26815, 28650, 
3 1 49 1 Yaqui Well: SDSNH 1 1 349 La Puerta: SDSNH 1 6998. 268 1 6 San Felipe Vallev; 
SDSNH 17009, 25632, 20849, 20850, 28755, 33022 The Narrows; SDSNH 3 1445 "5 
mi W of The Narrows: SDSNH 23442 Benson's Dry Lake; SDSNH 29078, 32993 
Borrego Valley: SDSNH 29110, 29514, 33398 Sentenac Canyon (vicinity); SDSNH 
33021, 33049, 33060 Scissors Crossing (vicinity). Riverside County: SDSNH 28670 
Cathedral City: SDSNH 31503 Indio; SDSNH 32137 Palm Springs R.R. Station. 

Remarks. — =Rhinochelus lecontei lecontei according to Shannon and Humphrey 
(1963, Herpetological 19(3):153-160). See also Medica (1975, SSAR Cat. Amer. Am- 
phib. Rept. 175.2). 

Salvadora grahamiae virgultea Bogert (1935) 

Bull. So. California Acad. Sci. 34(l):88-94 (p. 89). 

Holotvpe. — SDSNH 12025 a young adult male; Deerhorn Flat, San Diego Co., 
California'. By F. E. Walker, June 29, 1929. 

Remarks. — =Salvadora hexalepis virgultea according to Schmidt (1940, Zool. Ser. 
Field Mus. Nat. Hist. 24:143-150). 

Salvadora hexalepis klauberi Bogert (1945) 
Am. Mus. Novit. 1285:1-14 (p. 2). 

Holotvpe. — SDSNH 20912 an adult male; Cape San Lucas, Baja California, Mex- 
ico. By Fred Lewis, August 6, 1933 (preserved). 

Paratvpes.— W\ from Baja California as follows: SDSNH 3827 San Ignacio: SDSNH 
20466, 20511, 20858 Cape San Lucas; SDSNH 30385 3 mi S of Canipole, NW of 
Loreto; SDSNH 30386 5 mi N of San Xavier Mission; SDSNH 30387 Loreto. 

Salvadora hexalepis mojavensis Bogert (1945) 
Am. Mus. Novit. 1285:1-14 (p. 6). 

Paratvpes.— \s follows: Arizona: Mojave Co.: SDSNH 17315 Lucky Star Mine, 
Chemehuevis Mountains; SDSNH 25385 White Hills, 28 mi N of Chloride. Coconino 
Co.: SDSNH 34439 9 mi W of Cameron. California: Inyo Co.: SDSNH 25384 Towne's 
Pass, Panamint Mountains; SDSNH 28578 Linnie; SDSNH 32827 Daylight Pass, 
Funeral Mountains; SDSNH 34100 3 mi SW of Wildrose Station. Kern Co.: SDSNH 
25864 2 mi N of Grapevine Station; SDSNH 26 1 30 20 mi S of Inyokem. San Bernardino 
Co.: SDSNH 4400, 4401, 33446 Mountain Pass, Ivanpah Mountains; SDSNH 8503 
Twenty-nine Palms; SDSNH 10692 Klinefelter; SDSNH 33951 Hawes; SDSNH 35896 
7 mi W of Red Pass, NE of Barstow. Nevada: Clark Co.: SDSNH 25357 13 mi W of 
Indian Springs; SDSNH 25386 19 mi SE of Indian Springs near Com Creek. 

Remarks. — HoXoXy^Q-. AMNH 63000 an adult male. 

Sonora bancroftae Klauber (1943) 

Trans. San Diego Soc. Nat. Hist. 10(4):69-70 (p. 69). 

Holotvpe. — SDSNH 35077 a female; 2 mi E of San Jorge, Lower California, Mexico. 
By Mrs. Griffing [Margaret] Bancroft, April 10, 1942. 

Remarks. — "^Sonora semiannulata according to Frost (1983, Trans. Kansas Acad. 
Sci. 86(l):31-37). 

Sonora occipitalis klauberi Stickel (1941) 

Bull. Chicago Acad. Sci. 6(7): 135-140 (p. 138). 

Holotvpe. — SDSNH 29647 an adult male; Tucson, Pima Co., Arizona. By C. T. 
Vorhies, June 3. 1938. 

Paratype. — SDSNH 171 15 3 mi SE of Picacho, Pinal Co., Arizona. 

Remarks. — ^Chionactus occipitalis klauberi according to Stickel (1943, Proc. Biol. 
Soc. Wash. 56:109-128). 


Sonora palarostris KJauber (1937) 

Trans. San Diego Soc. Nat. Hist. 8(27):363-366 (p. 363). 

Holotype. — S>DSNY\ 26771 an adult male; 5 mi S of Magdalena, Sonora, Mexico. 
By George Lindsay, April, 1937. 

Remarks. —=Chionactis palarostris palarostris according to Klauber (1951, Trans. 
San Diego Soc. Nat. Hist. 1 1(9):141-204). 

Tantilla eiseni transmontana Klauber (1943) 

Trans. San Diego Soc. Nat. Hist. 10(5):71-74 (p. 71). 

Holotype. — SiySNW 29273 an adult male; 1 mi E of Yaqui Well, San Diego Co., 
California. By Charles Shaw and Cyrus Perkins, June 6, 1938. 

Paratvpes. — ¥vom California as follows: Riverside Co.: SDSNH 33760, Palm 
Springs. San Diego Co.: SDSNH 2633, 2634 Yaqui Well; SDSNH 11260 La Puerta; 
SDSNH 32419, 33997 Sentenac Canyon. 

Remarks. — = Tantilla planiceps transmontana according to Tanner (1966, Her- 
petologica 22(2): 134-1 52) and Tantilla planiceps according to Cole and Hardy (1981: 
268, Bull. Amer. Mus. Nat. Hist. 1 7 1 (3): 1 99-284), who regard the species as monotypic. 

Trimorphodon biscutatus semimtus Smith (1943) 

Proc. U.S. Natl. Mus. 93(3169):393-504 (p. 492). 

Paratype. — SDSNH 43401 10 mi N of Tafetan, Michoacan, Mexico. 

Remarks. — = Trimorphodon biscutatus biscutatus according to Duellman (1954, 
Occ. Pap. Mus. Zool. Univ. Michigan 560:1-24). Holotype: USNM 110410 a male 
(cited as EHT-HMS 23619 in Smith 1943). 


We are very grateful to James R. Dixon, Thomas H. Fritts, Darrel Frost, and Jay 
M. Savage for their reviews of earlier drafts. Marjorie Rea expertly typed the manuscript. 


Plethodon richmondi Netting and Mittleman (1938) 
Ann. Carnegie Mus. 27:287-293 (p. 288). 

Paratype.—SDSNW 32680 Oglebay Park near Wheeling, Ohio Co., West Virginia. 
Remarks. -Ho\oXypQ\ CM 14189 an adult male. SDSNH 32680 was formerly CM 


MOS. ^pp^^f?y SOCIETY OF 





Volume 20 Number 11 pp. 165-168 20 June 1984 

Imocaris tuberculata, n. gen., n. sp. (Crustacea: Decapoda) 
from the upper Mississippian Imo Formation, Arkansas 

Frederick R. Schram 

Department of Geology, San Diego Natural History Museum, San Diego, CA 92112 USA 

Royal H. Mapes 

Department of Geology, Ohio University, Athens, OH 45701 USA 

Abstract. A new genus and species of decapod crustacean is described from the Upper Missis- 
sippian Imo Formation, near Leslie, Arkansas. The exceptionally well-preserved specimen is a single 
carapace, associated with a moUuscan dominated fauna including ammonoids, gastropods, and bivalves. 
Imocaris tuberculata serves to help fill the stratigraphic gap between the oldest decapod, Palaeopalaemon 
newberryi in the Upper Devonian of Ohio and Iowa, and the diverse Triassic decapod faunas of Europe. 


Schram et al. (1978) redescribed Palaeopalaemon newberryi Whitfield, 1880, and 
recognized it as the earliest decapod crustacean. However, until now, there has been a 
significant gap in the decapod fossil record from this Upper Devonian form to the 
better known Triassic decapods of central Europe (see e.g., Forster, 1 967). The specimen 
described herein is a single well-preserved carapace from the Upper Mississipian- 
Chesterian (Naurian-Amsbergian; ammonoid zone E2b-c) of Arkansas interpreted to 
be a decapod crustacean. 

This specimen was collected from a road cut along Peyton Creek just south of the 
Van Buren Co. hne, Arkansas, along U.S. Hwy. 65 approximately 6.4 km (4 mi) 
southeast of Leslie, Arkansas. The fossil was found as the result of washing and con- 
centrating some 817 kgs of matrix from a 0.6 m (2 ft) concretionary shale layer ap- 
proximately 32 m (105 ft) above the contact of the Imo Formation with the underlying 
Pitkin Formation (in the middle of "bed 21," of Sutherland and Manger 1977, fig. 1). 

Systematic paleontology 

Order Decapoda Latreille, 1803 

Suborder Pleocyemata Burkenroad, 1963 

Infraorder Reptantia Boas, 1 800 

Section Brachyura Latreille, 1803 

Subsection Dromiacea de Haan, 1833 

Genus Imocaris n. gen. 

Z)/a^«05/5.— Carapace cylindrical, with subrectangular outline in both dorsal and 
lateral views. Surface tuberculate. Cervical and branchiocardiac grooves prominent. 
Rostrum small to non-existent. Antero-lateral margin with prominent denticles. 

Etymology.— ^3.mQd. after the Imo Formation, gender feminine. 

Type species. — Imocaris tuberculata n. sp. 










* . ' 


^^^jk: f ij-'*-''^', • 



':*f? ; ^''•''^^iv^-' 





 ' ---i- ..'. ■■,• ....-■• ' 




^^^^^K ' 

^^^^^C K 


y ^*BK - 

Figure 1. Imocaris tuberculata n. sp., holotype SDSNH 25139, stereo pair, scale = 5 mm. 

Imocaris tuberculata n. sp. 
Figure 1 

Diagnosis. — ^QcaM^t there is but one species, the diagnosis is the same as that of 
the genus. 

//o/orw?^.- SDSNH 25139 (Fig. 1). 

Loca//7y.- SDSNH loc. 3191; NE Va, sec. 11, T.13N., R.15W.; along road cut in 
U.S. Hwy. 65, south of Van Buren Co. line, Arkansas, on Peyton Creek. 

Stratum. — Imo Formation, Chesterian, Upper Mississippian. 

Etymology.— After the tuberculate nature of the carapace. 

Z)^^^/?//^^.— Anterior margin of carapace straight, without orbits, with only slight 
suggestion of rostrum. Lateral margin anterior of branchiocardiac groove with at least 
5 prominent denticles, and posterior of branchiocardiac groove marked by submarginal 
furrow. Posterior margins slightly concave, with faint submarginal furrow. Prominent, 
deep cervical and branchiocardiac grooves, cervical grooves extend laterally from mid- 
line and turn anteriad as short antennar grooves, branchiocardiac grooves continuous 
laterally with prominent inferior grooves. Carapace surface tuberculate. Mid-dorsal line 
marked by row of 4 large tubercles between rostral area and cervical groove, by slight 
groove between the cervical and branchiocardiac grooves, and by clear ridge sur- 
mounted by row of 5 (?6) tubercles between branchiocardiac groove and posterior 
margin. Pair of tubercle rows located laterally between branchiocardiac groove and 
posterior margin about '/2-way between median ridge and lateral margins, forming 
planes at which curvature of posterior carapace surface changes from horizontal to 
more vertical orientation. Carapace mid-line length— 1 1 mm, approximate lateral mar- 
gin length— 12 mm, width — 9.5 mm. 

Remarks. — The fossil preserves an excellent view of the dorsal surface of the 
carapace. Attempts at preparation of the underside and margins of the specimen, 
however, seem to indicate that only the carapace is fossilized, i.e., no ventral thoracic 
structures appear to be present. 



The higher taxonomic affinity of any Paleozoic decapod is of interest because of 
the understanding it might lend to elucidating the sequence of events in the early 
radiation of Decapoda. Paleozoic decapods have been rare and widely scattered geo- 
graphically. The type specimens of the supposed Permian decapod carapaces from 
Sicily, Palaeopemphix Gemmellaro, 1890, have been lost and, thus, their suggested 
decapod versus cumacean affinities can never be verified. The Devonian Palaeopalae- 
mon newberryi, preserved in great detail, possesses several features in a combination 
of both astacidean and palinuran reptants (Schram et al. 1978); and, in addition, 
Felgenhauer and Abele (1983) have noted a characteristic natatian feature in that form 
as well, viz., the large scaphocerite. Bachmeyer and Malzahn (1983) have recorded 
from the Upper Permian Zechstein of Germany a free decapod cheliped (given the 
name Erymastacus ? hoerstgenensis), as well as leg and carapace fragments attributed 
to decapods in the stomach contents of the chimaeriform fish Janassa hituminosa. The 
fragmentary nature of the Zechstein fossils, however, makes them difficult to compare 
to the other known Paleozoic decapod whole body fossils. Finally, little can be said 
about the Soviet Permian form Protoclytiopsis antiqua Birshtein, 1958, except that it 
appears to be taxonomically close to a large, reptant, glypheoid type (Schram 1980). 

Several characters mark Imocahs tuberculata as unusual, viz., the cylindrical form 
and rectangular proportions of the carapace, the presence of only a branchiocardiac 
groove posterior to the cervical (or alternatively a fusion of the post-cervical and 
branchiocardiac grooves), the lack of orbits, and the small or absent rostrum. Unfor- 
tunately, the lack of preservation of any eyes, antennae, mouthparts, thoracopods, and 
abdomen makes a definitive higher taxonomic assignment difficult. 

Quite unexpectedly, the closest analogs to the derived features of Imocaris tuber- 
culata listed above are to be found among the dromiacean brachyurans. These living 
forms tend to possess markedly cylindrical and somewhat subrectangular carapaces, 
strong cervical and branchiocardiac grooves, absent or poorly developed orbits, fre- 
quently weakly developed rostra, and (in at least the prosopids, some dynomenids, and 
the homoloids) prominent carapace sculpturing. The assignment of Imocahs to the 
dromiaceans on the basis of this single specimen considerably extends the range of that 
taxon back from the Lower Jurassic into the middle of the Carboniferous. 

The recognition of Imocaris as a decapod begins to fulfill the expectations of Schram 
et al. (1978) that an extensive Paleozoic radiation of Decapoda is to be discovered in 
upper Paleozoic strata. 


This research is supported in part by NSF grant BSR 82-12335 (FRS) and a grant 
from the Ohio University Research Council (RHM). 

Literature Cited 

Bachmeyer, F., and E. Malzahn. 1983. Der erste 
Nachweis eines decapoden Krebses im nie- 
derrheinischen Kupferschiefer. Annalen des 
Naturhistorischen Museums in Wien. 85/A:99- 

Birshtein, Ya. A. 1958. Drevneishii predsta vitel 
otryada desyatinogikh rakoobraznikh, Proto- 
clytiopsis antiqua, in Permakikh otiozhenii za- 
padnoi Sibiri. Doklady Akademii Nauk SSSR 

Felgenhauer, B. E., and L. G. Abele. 1983. Phy- 
logenetic relationships among shrimp-like 
decapods. Crustacean Issues 1:291-31 1. 

Forster, R. 1967. Die reptanten Dekapoden der 

Trias. Neues Jahrbuch fur Geologic und Pa- 
leontologie Abhandhungen 128:136-194. 

Gemmellaro, G. G. 1890. Crostacei dei calcari 
con Fusulina della Valle del Fiume Sosio, nella 
provicia di Palermo in Sicilia. Societa Italiana 
di Scienze Naturali, Mcmorie (3)8:1-40. 

Schram, F. R. 1980. Notes on miscellaneous crus- 
taceans from the Late Paleozoic of the Soviet 
Union. Journal of Paleontology 54:542-547. 

, R. M. Feldmann, and M. J. Copcland. 1978. 

The Late Devonian Palaeopalaemonidae and 
the earliest decapod crustaceans. Journal of Pa- 
leontology 52:1375-1387. 

Sutherland, P. K., and W. L. Manger. 1977. Up- 
per Chesterian-Morrowan Stratigraphy and the 


Mississippian-Pennsylvanian Boundary in Whitfield, R. P. 1880. Notice of new forms o. 

Northeastern Oklahoma and Northwestern fossil crustaceans from the Upper Devonian 

Arkansas. Oklahoma Geological Survey rocks of Ohio, with description of new genera 

Guidebook 1 8. University of Oklahoma, Nor- and species. American Journal of Science (3)19: 

man. 33-42. » 





Volume'^ Number 12 pp. 169-188 20 November 1984 

New material of Hydrodamalis cuestae (Mammalia: Dugongidae) from 
the Miocene and Pliocene of San Diego County, California 

Daryl P. Domning 

Department of Anatomy, Howard University. Washington, D.C. 20059 

Thomas A. Demere 

Department of Geology, Natural History Museum, P.O. Box J 390, San Diego, CA 92112 

Abstract. The geology, faunal content, and age of sirenian-bearing marine rocks in San Diego 
County, southern California, are reviewed and reevaluated. The San Mateo Formation comprises two 
members, respectively of Late Miocene (late Clarendonian or, more likely, early Hemphillian) and 
Early Pliocene (late Hemphillian) age. The San Diego Formation also includes two members, both 
probably of Late Pliocene (Blancan) age, though the upper member may extend into the Early Pleis- 
tocene. All four members have yielded new specimens of sirenians, most or all of which represent 
Hydrodamalis cuestae Domning 1978. These specimens extend our knowledge of the osteology of this 
species, and confirm the supposition that the holotype is abnormal in several respects. During the 
Hemphillian, the juveniles of the species had an upper dentition probably consisting of DP^-^; it is still 
unknown whether later Pliocene juveniles retained teeth. A braincase from the San Diego Formation 
is the largest known of any sirenian, probably representing an individual over 10 m long and confirming 
the previous observation that Hydrodamalis grew larger in California than at Bering Island. 


Domning ( 1 978) reviewed the fossil record of North Pacific sirenians and described 
the new species Hydrodamalis cuestae, a form intermediate between the Late Miocene 
Dusisiren jordani (Kellogg 1925) and the Late Pleistocene-Recent H. gigas (Zimmer- 
mann 1780) (Steller's sea cow). The holotype of H. cuestae came from the Upper 
Pliocene (Blancan) Pismo Formation in San Luis Obispo County, California, and re- 
ferred specimens came from the Hemphillian San Mateo Formation in San Diego 
County, the Hemphillian Capistrano Formation in Orange County, and an unnamed 
Pliocene unit in Baja California, Mexico. A single thoracic vertebra from the Blancan 
San Diego Formation near Tijuana, Mexico, was identified only as Hydrodamalis 

Since 1978, a number of additional specimens o^ Hydrodamalis cuestae have been 
obtained by the San Diego Natural History Museum from the San Mateo and San 
Diego Formations in San Diego County. These clarify some of the problems posed by 
the previously known material; in particular, they show that certain anomalous features 
of the holotype, thought by Domning (1978) to be pathological, are indeed atypical of 
the species. 

Abbreviations used are as follows: SDSNH, San Diego Society of Natural History; 
UCMP, University of California Museum of Paleontology, Berkeley; USNM, U.S. 
National Museum of Natural History, Washington, D.C. 


Geology and Age of the Deposits 

Remains of fossil hydrodamalines from San Diego County, California, have been 
recovered from both the Upper Miocene-Lower Pliocene San Mateo Formation (Hemp- 
hillian) at Oceanside and the Upper Pliocene San Diego Formation (Blanc^n) at San 
Diego. These rock units accumulated in separate sedimentary basins and contain dis- 
tinctly different marine vertebrate assemblages. 

San Mateo Formation 

Barnes (1976) and Barnes et al. (1981) have tentatively assigned the vertebrate- 
producing marine beds at Oceanside to the San Mateo Formation of Woodford (1925). 
This rather poorly defined rock unit has been mapped from the type area near San 
Clemente throughout the coastal portion of Camp Pendleton Marine Corps Base as far 
south as the city of Oceanside (Moyle 1973, Young and Berry 1981). The best exposures 
of the San Mateo Formation at Oceanside are in the Lawrence Canyon area adjacent 
to the San Luis Rey River (Fig. 1 ). Here erosion, grading, and quarry operations have 
combined to produce a number of natural and artificial outcrops. 

Lawrence Canyon is aligned in part along a north-south striking, eastward-dipping, 
high angle normal fault. The San Mateo Formation is confined to the east side of this 
fault where it rests unconformably on westward-dipping strata of the Middle Miocene 
San Onofre Breccia and in turn is overlain unconformably by flat-lying Upper Pleis- 
tocene, nonmarine terrace deposits. In Lawrence Canyon the San Mateo Formation is 
approximately 24 m thick and, as noted by Barnes et al. (198 1), is divisible into a lower 
and an upper unit. The lower unit consists of white, fine-grained, massive, friable, 
micaceous sandstones with occasional green or black claystone lenses and locally com- 
mon pebbles and cobbles. The upper unit is a complexly bedded sequence of gravels, 
pebble to cobble conglomerates and friable sandstones. A sharp unconformity marked 
in places by scour and fill features separates the upper gravel conglomerate unit from 
the lower white sandstones. Marine vertebrate fossils have been collected from both 
the upper and the lower unit (Barnes 1976, Domning 1978, Barnes et al. 1981, Howard 
1982) whereas invertebrate fossils are conspicuously absent. Barnes et al. (1981) have 
assigned the vertebrate fossil assemblages from the lower and upper units to the San 
Luis Rey River Local Fauna and the Lawrence Canyon Local Fauna, resnectively. 

Fossils in the lower white sandstones generally occur as single isolated elements 
and occasionally as associated partial skeletons. In contrast, fossils from the upper 
gravel conglomerate unit are often concentrated in distinct "bone horizons." One par- 
ticular horizon (SDSNH locality 3161) is a 1.2 m thick, fining-upward sequence 
divisible into four lithologic units: 1) a basal clast-supported pebble to cobble con- 
glomerate; 2) a matrix-supported pebble conglomerate; 3) an interval of poorly bedded 
sandstones and gravels with laminated fine-grained sandstone lenses; and 4) thickly 
laminated fine- and coarse-grained sandstones. Fossils are concentrated in the matrix- 
supported pebble conglomerate and generally occur as broken and abraded fragments 
of large bones or as complete and differentially preserved smaller resistant elements. 
Particularly common are shark teeth, cetacean earbones, and bird humeri and ulnae. 
Larger bones generally occur in association with the larger pebbles or cobbles, suggesting 
size-sorting by currents. Apparently the fossils were transported and deposited as bio- 
genic clasts within the traction load. 

In terms of general depositional environments it appears that the lower white 
sandstone unit was deposited under "normal" marine conditions, perhaps at middle 
to inner shelf depths. The dominance of sandstones over finer-grained lithologies would 
support this idea of shallow water deposition, although without a preserved benthonic 
assemblage (e.g., mollusks) it is not possible to make any definite paleobathymetry 
estimates. Clearly, however, because of the rich marine vertebrate assemblage, this is 
a marine unit. 

The upper gravel conglomerate unit, in contrast to the lower white sandstones, is 
characterized by complex bedding and very coarse-grained lithologies. The bedding 


33 00 

5 m I 

10 km 

— 32 30 

I 1 1 7 00 

Figure 1. Index map of western San Diego County showing the principal outcrop areas of rocks containing 
Hydrodamalis cuestae. 1, Hemphillian San Mateo Formation, Lawrence Canyon; 2, Blancan San Diego 
Formation, northern San Diego Mesa; 3, Blancan San Diego Formation, Chula Vista/National City area. 

features are interpreted as representing anastomosing channel-fill deposits and, together 
with the coarse-grained lithologies, suggest a fluvial environment. However, the oc- 
currence of common marine vertebrate fossils (not reworked from the lower unit) points 
instead to a marine environment. The model herein proposed to accommodate these 


apparently conflicting environmental settings involves deposition of the upper gravel 
conglomerate unit at the distal, submarine margin of a river-dominated gravel-cobble 

Fossil remains of Hydwdamalis cuestae occur in both the San Luis Rey River 
Local Fauna (lower white sandstone unit) and the Lawrence Canyon Local Fauna (upper 
gravel conglomerate unit). Both local faunas contain a variety of taxa including sharks, 
rays, bony fishes, sea birds, fur seals, walrus, and toothed and baleen whales (Barnes 
et al. 1981). Recent work by personnel at the San Diego Natural History Museum has 
added additional taxa, especially species of birds (R. M. Chandler, personal commu- 
nication), to the published faunal lists of Barnes et al. (1981) and Howard (1982). To 
the San Luis Rey River Loca Fauna can be added the white shark Carcharodon me- 
galodon Agassiz, a new large species of the flightless auk Praemancalla, a shearwater 
(Puffinus species), an eagle (Accipitriformes), a mysticete (Balaenopteridae), and the 
camel Aepycamelus species (S. D. Webb, personal communication). To the Lawrence 
Canyon Local Fauna can be added a billfish (Istiophoridae), a cormorant (Phalacrocorax 
species), a shearwater {Puffinus species), the loon Gavia concinna Wetmore, an albatross 
(Diomedea species), a mysticete (Balaenopteridae), a llama (cf Hemiauchenia species; 
S. D. Webb, personal communication), and a peccary (Tayassuidae). These last two 
taxa and the camel {Aepycamelus species) are part of a small terrestrial mammal as- 
semblage from the San Mateo Formation which also includes horses of the genus 
Dinohippus (or possibly Pliohippus; B. J. MacFadden, personal communication). 

Barnes et al. (1981) have used the joint occurrence of the horse (Pliohippus or 
Dinohippus) and the sabertoothed salmon Smilodonichthyes rastrosus Cavender and 
Miller to correlate both the upper and lower unit of the San Mateo Formation at 
Oceanside with the Hemphillian North American Land Mammal Age. The recent 
recovery of an isolated M- belonging to the camelid Aepycamelus species from the 
lower white sandstone unit offers a refinement of this correlation and suggests that the 
San Luis Rey River Local Fauna is of late Clarendonian or early Hemphillian age 
(Webbet al. 1981). 

As suggested by Howard (1982), the avifauna of the San Mateo Formation (es- 
pecially the mancalline taxa) may offer a means for finer biostratigraphic resolution in 
the section. The occurrence of the genus Praemancalla in the lower unit and the genus 
Mancalla in the upper unit is the basis for this biostratigraphy. Previously, the genus 
Praemancalla was known only from Clarendonian-aged rocks (Monterey Formation) 
in Orange County (Howard 1976), with two species assigned to this genus. The older 
species P. lagunensis Howard is recorded from the lower part of the Monterey For- 
mation in Orange County, while P. wetmorei Howard is recorded from the upper part 
of the formation. Howard (1982) referred material from the lower white sandstone unit 
at Oceanside to P. species cf P. wetmorei, noting that this was the first Hemphillian 
record of Praemancalla. In the upper gravel conglomerate unit Praemancalla is replaced 
by the genus Mancalla, represented here by three species, M. cedrosensis Howard, M. 
milleri Howard, and M. diegensis (Miller). The co-occurrence of these three species 
appears to define a useful assemblage zone. Howard (1982:13) has suggested that a 
considerable hiatus is represented by the unconformity within the San Mateo For- 
mation, one that allowed ". . . time for the demise of the Praemancalla and the evolution 
of Mancalla.'^ It seems then, that the range of Praemancalla defines a biostratigraphic 
interval and that the hiatus in the San Mateo Formation may be represented elsewhere 
by an additional and younger biozone. From discussions with R. M. Chandler {personal 
communication) it seems that such a biozone is preserved in mudstones of the Cap- 
istrano Formation at San Clemente. Here an avifauna was recovered which contains 
Mancalla californiensis Lucas, M. cedrosensis, and the auklet Cerorhinca minor How- 
ard. (Unfortunately it has not been possible to place the stratigraphic position of this 
fossil locality within a composite section for the very thick and long-ranging Capistrano 
Formation.) Apparently this assemblage zone is also preserved within the basal portion 
of the Almejas Formation on Cedros Island, Baja California, Mexico, where Howard 
(1971) recorded M. cedrosensis and C m/nor (although without M. californiensis). 


To summarize these assemblage zones, we then have: Praemancalla lagunensis in 
the Monterey Formation (Clarendonian); P. wetmorei in the Monterey Formation 
(Clarendonian) and "lower" San Mateo Formation (late Clarendonian or early Hemp- 
hillian); Mancalla cedrosensis and Cerorhinca minor with or without M. californiensis 
in the Upper Miocene portions of the Almejas Formation (Hemphillian) and Capistrano 
Formation (Hemphillian); M. cedrosensis, M. milleri, and M. diegensis in the "upper" 
San Mateo Formation (late Hemphillian); and to complete the zonation. M. milleri, 
M. diegensis, and M. emlongi Olson in the San Diego Formation (Blancan). (These 
correlations and age assignments are based in part on the work of Barnes 1976, Re- 
penning and Tedford 1977, Howard 1978, 1982, and R. M. Chandler, personal com- 
munication.) Whether the "lower" San Mateo Formation is late Clarendonian or early 
Hemphillian in age cannot at this time be resolved biostratigraphically. However, 
physical stratigraphy tends to support an early Hemphillian age assignment. Vedder 
(1972) has suggested that the San Mateo Formation (at least in the type area) may be 
a channel facies within the lower part of the Capistrano Formation. Similarly, Ehlig 
(1979) has mapped the San Mateo Formation near San Onofre as a member of the 
Capistrano Formation. Although the nomenclatural standing of the San Mateo For- 
mation is questionable (it is retained here pending completion of ongoing field work 
at Camp Pendleton), its correlation with the Hemphillian Capistrano Formation (and 
not the Clarendonian-aged Monterey Formation) appears to be certain. Thus the lower 
white sandstones at Oceanside probably are correlative with the basal portion of the 
Capistrano Formation (Upper Miocene, early Hemphillian), while the upper gravel 
conglomerate unit at Oceanside probably correlates with some horizon near the top of 
the Capistrano Formation (Lower Pliocene, late Hemphillian). Howard (1982) has 
proposed a similar Late Miocene-Early Pliocene correlation for the San Mateo For- 
mation at Oceanside, in contrast to the strictly Miocene age assignment of Barnes et 
al. (1981). 

San Diego Formation 

The geology of the marine Pliocene San Diego Formation has recently been sum- 
marized by Demere ( 1 983). This rock unit extends in a broad area of outcrop throughout 
much of the southwestern portion of San Diego County (Fig. 1). The formation is 
fossiliferous throughout this area of outcrop and contains a diverse and well-preserved 
assemblage of both marine invertebrate and vertebrate taxa. 

The San Diego Formation was deposited during a marine transgression of the 
Neogene San Diego Basin, which like other onshore sedimentary basins in southern 
California (e.g., Ventura Basin, Los Angeles Basin) is structurally related to the wrench 
and extensional tectonics of the continental borderland. Deposition began during the 
Late Pliocene and possibly continued into Early Pleistocene time, accumulating at least 
75 m of marine and 9 m of nonmarine sedimentary rocks. The overall stratigraphic 
sequence suggests a successive filling and shallowing of this basin. It is now apparent 
that extensional tectonics have controlled both the initial deposition as well as the 
present outcrop distribution of this rock unit. Numerous high-angle normal faults 
striking north to northwest cut the area into a series of fault blocks which expose 
different portions of the Pliocene section. 

In an attempt to correlate these various fault blocks. Demere ( 1 983) has informally 
subdivided the San Diego Formation into a "lower" and an "upper" member using 
both lithologic and paleontologic criteria. The "lower" member is characterized by 
yellowish, very fine-grained, massive, friable, micaceous sandstones with locally well- 
bedded sequences of laminated and cross-bedded sandstones, pebble to cobble con- 
glomerates and well-cemented shell beds. This "lower" member is richly fossiliferous 
and has produced the bulk of the marine invertebrate fauna so well known through 
the work of Grant and Gale (1931) and Hertlein and Grant (1944, 1960, 1972). In 
addition, the diverse avifaunas (Howard 1949, Miller 1956) and cetacean assemblages 


(Barnes 1973, 1976) reported from the San Diego Formation have been largely collected 
from the "lower" member. 

Lithologically the "upper" member is characterized by well-bedded sequences of 
pebble to cobble conglomerate, well-cemented fossiliferous sandstones, and^medium- 
to coarse-grained friable sandstones. Marine invertebrate fossils are locally common 
in this member, which to date has produced only a few vertebrate remains. 

Vertebrate fossils in the San Diego Formation generally occur as single isolated 
skeletal elements, although occasionally partial or complete skeletons are found. A few 
rare "bone beds" containing concentrated, unassociated elements have recently been 
discovered in the formation. To date, the most productive fossil sites have been in the 
Mission Hills area near downtown San Diego and in the eastern portions of National 
City and Chula Vista. In these areas large-scale grading operations have provided very 
extensive exposures of the San Diego Formation, although in most cases these new 
exposures are accessible for only a short period of time because of development and 

Found in association with the vertebrates are rich assemblages of marine macroin- 
vertebrates, primarily mollusks, which provide both biostratigraphic and paleoenvi- 
ronmental control. The "lower" member contains a middle to outer shelf molluscan 
fauna characterized by Patinopecten healeyi (Arnold), Pecten stearnsii Dall, Lucinoma 
annulata (Reeve), and Opalia varicostata (Steams). In contrast, mollusks from the 
"upper" member indicate deposition in littoral to inner shelf depths. Characteristic 
species include Pecten bellus (Conrad), Argopecten hakei (Hertlein), and Nucella la- 
mellosa Gmelin along with the echinoid Dendraster ashleyi (Arnold). Both members 
reflect normal marine deposition in a broad coastal embayment probably similar to 
present-day Monterey Bay along the central California coast. 

Fossil remains of Hydrodamalis cuestae have been recovered from both the "low- 
er" and "upper" members of the San Diego Formation. As presently understood, the 
aggregate vertebrate faunule from the "lower" member consists of 7 species of sharks 
and rays, more than 50 species of bony fishes (most of which are known only from 
otoliths), 24 species of sea birds, an otariid and a dusignathine (odobenid) pinniped, 
7 odontocetes, 10 mysticetes, and the sirenian. A few terrestrial mammal taxa including 
horse {Equus species), camel (cf. Titanotylopus species), peccary (cf. Platygonus species; 
M. O. Woodbume, personal communication), and gomphothere {Stegomastodon cf. S. 
rexroadensis Woodbume) occur in association with the diverse marine vertebrate as- 
semblage. The "upper" member has produced only three vertebrate taxa to date: an 
albatross {Diomedea species), a mysticete (cf. Balaenopteridae), and the sirenian. 

Bames (1976) has correlated the San Diego Formation ("lower" member) with the 
Blancan North American Land Mammal Age. This correlation, based on the occurrence 
of the horse Equus, is supported by the recent discovery of teeth referable to the 
gomphothere Stegomastodon cf. S. rexroadensis (M. O. Woodbume, personal com- 
munication). Relying on the stratigraphic ranges of molluscan species, the "lower" 
member is correlative with the San Joaquin Formation in the San Joaquin Basin, the 
Careaga Formation in the Santa Maria Basin, and the Niguel Formation and the upper 
Femando Formation in the southeastem Los Angeles Basin, all Late Pliocene in age 
(Woodring and Bramlette 1950, Vedder 1972). In tum, the "upper" member of the 
San Diego Formation correlates with the lower part of the Santa Barbara Formation 
in the Ventura Basin, which is considered to be Late Pliocene to Early Pleistocene in 
age (Keen and Bentson 1944). The meager microfossil evidence available (Ingle 1967, 
Mandel 1973) suggests that the San Diego Formation is apparently no older than 
planktonic foraminiferal zone N.21 (approximately 3.0 million years B.P., Late Plio- 
cene) and is perhaps as young as the Emiliania annula calcareous nannoplankton 
subzone (approximately 1.5 million years B.P., Early Pleistocene; E. D. Milow, personal 
communication). This Pleistocene correlation is tenuous at the moment and must await 
completion of additional field work for confirmation. It should not be interpreted to 
mean that the entire "upper" San Diego Formation is Pleistocene. 



Order Sirenia 

Family Dugongidae 

Subfamily Hydrodamalinae 

Hydrodamalis cuestae Domning, 1978 


San Mateo Formation 

San Luis Rev River Local Fauna (of Barnes et al. 1981).— 
SDSNH locality 2957 (=UCMP loc. V68144), Loretta St., Oceanside: 

SDSNH 22655: Two rib fragments. Coll. J. W. Tobiska, I 1979. 
SDSNH locality 3004 (=UCMP loc. V68145), Loretta St., Oceanside: 

SDSNH 21076: Juvenile left maxilla. Coll. T. A. Demere, 15 III 1980. 
SDSNH locality 3134 (=UCMP loc. V68147), Lawrence Canyon, Oceanside: 

SDSNH 23384: Proximal end of rib. Coll. R. A. Cerutti, 22 III 1981. 

Lawrence Canyon Local Fauna (of Barnes et al. 1981).— 
SDSNH locality 3161 (=UCMP loc. V68106), Lawrence Canyon, Oceanside: 
SDSNH 24413 

SDSNH 24454 
SDSNH 24685 

Fragment of juvenile left maxilla. Coll. SDSNH party, VII 1982. 
Rib fragment. Coll. SDSNH party, VII 1982. 
Immature right humerus. Coll. B. O. Riney and R. A. Cerutti, 26 
VI 1982. 
SDSNH 24686: Proximal end of juvenile ?right ulna. Coll. SDSNH party, VII 


San Diego Formation 

' 'Lo wer ' ' Mem ber. — 

SDSNH locality 2970-B, Washington St., San Diego: 

SDSNH 21685: Rib fragment. Coll. R. A. Cerutti, IX 1980. 

SDSNH locality 3148, California St., San Diego: 

SDSNH 23719: Vertebrae CI -2, C6-7, Tl-?6. Coll. R. A. Cerutti and T. A. De- 
mere, 16 II 1981. 

SDSNH locality 3172, Florida Canyon, San Diego: 

SDSNH 24679: Two immature thoracic vertebrae. Coll. R. A. Cerutti and T. A. 

Demere, 2 XII 1981. 

SDSNH locality 3 1 74, H Street extension, Chula Vista: 

SDSNH 24683: AduU right rib. Coll. R. A. Cerutti and B. O. Riney, 24 VIII 1982. 

SDSNH locality 3175, Adams Ave., San Diego: 

SDSNH 24684: Anterior end of immature left mandible. Coll. R. H. Norwood, 

SDSNH 24687: Rib fragment. Coll. R. H. Norwood, 1975. 

SDSNH locality 3158, Hidden Vista, Chula Vista: 

SDSNH 23726: Adult braincase. Coll. R. A. Cerutti, 15 IX 1981. 


SDSNH 2468 1 : Distal ends of immature ?right radius and ulna. Coll. R. A. Cerutti, 

20 VII 1981. 
SDSNH 24682: Seven rib fragments. Coll. R. A. Cerutti, 22 VII 1981. 

"Upper" Member. — 

SDSNH locality 3173, 38th and Beech Streets, San Diego: 

SDSNH 24680: Neural arch of immature thoracic vertebra {Hvdrodamalis species 

indet.). Coll. T. A. Demere, 21 I 1983. 

Description of Specimens from the San Mateo Formation 

A/axz7/a. — A juvenile left maxilla (SDSNH 21076, Fig. 2a-e), as noted by Barnes 
et al. (1981), is almost identical to the right maxilla (UCMP 86345) previously reported 
from the same locality (Domning 1978: tab. 24; pi. 17, fig. a). It is exactly the same 
size and represents the same growth stage, but is more complete, measuring 129 mm 
in overall length and 58 mm in height. (It may even represent the same individual.) 
The first alveolus, for a single-rooted tooth, lies slightly posterior to the zygomatic- 
orbital bridge and is 7.5 mm long. The second alveolus, for a two-rooted tooth, is 8.5 
mm long, and the third, for a three-rooted tooth with apparently coalesced roots, is 
about 9 mm long and 7 mm wide. The combined length of these three alveoli is 27 
mm. Posterior and dorsal to the third alveolus is a large broken dental capsule (1 1 mm 
in dorsoventral height) for an unerupted tooth. The narrowest part of the palatal surface 
(20 mm wide, left half only) lies just forward of the zygomatic-orbital bridge, and 
consists of a flat surface bordered medially by a shallow palatal gutter (containing a 
foramen anteriorly and ending posteriorly in another foramen) and laterally by a sharp 
edge where it meets the lateral surface perpendicularly. The zygomatic-orbital bridge 
is elevated 19 mm above the palatal surface, and is 30 mm long anteroposteriorly and 
10 mm thick. The vertical plate of the bridge, which articulated with the jugal, is 55 
mm in height and contacts the main body of the maxilla above the infraorbital foramen; 
the latter is invisible in ventral view. On the ventromedial wall of the foramen are the 
anterior and posterior openings of a short, horizontal canal (a continuation of the 
premaxillary canal); medial to the posterior opening is a larger pit or canal opening 
posteriorly. The palate is a maximum of 19 mm thick. 

A fragmentary left maxilla of a larger juvenile (SDSNH 2441 3) has a palatal surface 
which slopes dorsolaterad just forward of the zygomatic-orbital bridge, as in more 
mature individuals of Hydrodamalis. At the level of the front side of the bridge, this 
surface is only 18 mm wide, proportionately narrower than in the above specimen. 
The lateral surface bears a prominent groove (continuation of the premaxillary canal) 
anterior to the infraorbital foramen. The palate reaches a thickness of 31 mm, and the 
intermaxillary suture bears numerous deep vertical interdigitations. No dental alveoli 
are preserved. 

Dentition. — TttXh o^ Hydrodamalis cuestae are still unknown, but the essentially 
complete maxilla described above (SDSNH 21076) allows us to speculate on the ho- 
mologies of the teeth present in the juvenile. The pattern of roots of the three fully 
erupted teeth matches that of the teeth identified as DP-"" in Dusisiren (Domning 1 978); 
this implies that the dental capsule contained DP^ In view of the immediate descent 
of H. cuestae from Dusisiren (Domning 1978), this seems the most parsimonious 
interpretation. However, in living Dugong (Marsh 1980) and Trichechus (Domning 
1 982), the three most anterior cheek teeth of the juvenile appear to be the only premolars 
(DP'-5). A fuller growth series of//, cuestae is needed to settle this question conclusively, 
and also to determine whether Blancan as well as Hemphillian juveniles of the species 
still possessed teeth. 

Humerus.— \n immature right humerus (SDSNH 24685, Fig. 5) is complete except 
for the unfused epiphyses. In size, it falls between the two immature humeri of Hy- 
drodamalis gi gas described by Whitmore and Gard ( 1 977); in shape, it closely resembles 


Figure 2. Hydrodamalis cuestae. a-e, juvenile left maxilla. SDSNH 21076. a, lateral view; b, medial view, 
c, dorsal view; d, ventral view; e, posterior view, f-h, atlas. SDSNH 23719. f, dorsal view; g, anterior view; 
h, posterior view. Scales = 4 cm. 


Table 1. Skull measurements of Hydrodamalis cuestae (SDSNH 23726) from the San Diego Formation, 
in millimeters. Letters in parentheses denote measurements used by Domning (1978: tab. 2, fig. 7); e = 

Rear of occipital condyles to anterior end of interfrontal suture (BI) » 4 1 8e 

Length of interfrontal suture 143e 

Length of skull roof to rear of external occipital protuberance 310 + 

Top of supraoccipital to ventral side of occipital condyle (de) 223 

Breadth across occipital condyles (fF) 276e 

Width of supraoccipital 250 

Height of supraoccipital to top of external occipital protuberance 120e 

Breadth of cranium at frontoparietal suture (GC) 123 

Width of foramen magnum (gg') 125e 

Height of foramen magnum (hi) 64 

Minimum width of basioccipital 78 

Length of right zygomatic process of squamosal (O'P') 229 + 

Anterior tip of zygomatic process to rear edge of squamosal below mastoid foramen (O'T') 293 + 

Frontoparietal suture to rear of external occipital protuberance (P) 169e 

Anteroposterior length of root of zygomatic process (Q'R') 103 

Length of cranial portion of squamosal (S'T') 196 + 

Posterior height of cranial portion of squamosal (U'V) 182 

Dorsoventral breadth of zygomatic process (W'X') 1 17 

the smaller of these (USNM 186807), differing from the other (USNM 170761) in 
lacking a pronounced "shoulder" above the ectepicondyle. The deltoid crest is massive 
and rounded, without a recurved flange. The total length (without epiphyses) is 291 
mm; the proximal breadth, 130 mm; and the distal breadth, 156 mm. 

Ulna.— T\iQ proximal end of a juvenile ?right ulna (SDSNH 24686) has a width 
of 65 mm; the shaft is thicker anteroposteriorly than mediolaterally. It compares well 
in proportions with the ulna of a much larger immature H. gigas (USNM 170761) 
from Amchitka (Whitmore and Gard 1977:pl. 8, figs. 3-5). 

Ribs.—ThQ proximal end of a subaduh right rib (SDSNH 22655) is very flat (65 x 
34 mm) and has a tiny tubercle whose lateral edge is 74 mm from the tip of the 
capitulum. The distal end of an immature rib (SDSNH 24454) measures 59 x 54 mm 
in diameter and has a broad, concave, rugose distal surface for cartilage attachment. 
A zone of cancellous bone is partly exposed by bone resorption on its medial side. The 
proximal end of a larger, more posterior left rib (SDSNH 23384) has the capitulum 
and tubercle coalesced. Both it and SDSNH 22655 are completely dense where broken. 

Description of Specimens from the San Diego Formation 

Skull.— A nearly complete adult braincase (SDSNH 23726, Fig. 6, Table 1), in- 
cluding the right zygomatic process of the squamosal but lacking the pterygoid processes, 
expands our knowledge of the occipital region of Hydrodamalis cuestae. The interior, 
however, is still filled with very hard matrix, precluding examination of the internal 
structures. The dimensions of the preserved portions indicate that this individual was 
larger than any other Hydrodamalis or other sirenian ever found, and in life no doubt 
measured well over 10 m in body length. 

Frontal: The supraorbital processes are missing, but the anterior border seems to 
be complete. It is arched upward somewhat at the midline, due to a broad, prominent 
median boss on the dorsal surface. Lateral to this are depressions bordered by upraised 
forward extensions of the temporal crests. The lateral surfaces drop almost perpendic- 
ularly from these crests, with little or no overhang. 

Parietal: The cranial vault is square anteriorly with no overhang of the temporal 
crests, which are nearly parallel anteriorly but posteriorly form sharp ridges which 
diverge and descend to meet the dorsal ends of the sigmoid ridges of the squamosals. 
There is only a slight indentation in the dorsal surface where it meets the top of the 


squamosal, much as in H. gigas. The parietal roof is slightly concave anteriorly and 
flat posteriorly. 

Supraoccipital: The width/height ratio is 2.08, being within the previously reported 
range of variation. The supraoccipital forms an angle of 124° with the after part of the 
parietal roof and is within the range of variation of the USNM sample of Bering Island 
H. gigas. The external occipital protuberance is low and indistinct, with no median 
ridge extending below it. The lateral borders are rounded, rugose, and extremely thick 
and massive. 

Exoccipital: The sutures with the supraoccipital are indistinct, but the exoccipitals 
apparently fail to meet in the dorsal midline. The dorsal border of the foramen magnum 
is very gently arched with almost no dorsal peak, as in H. gigas. The dorsolateral border 
of the exoccipital is very thick, rounded, and rugose, and overhangs posteriorly, but is 
not expanded to the degree seen in H. gigas. However, it does confirm the supposition 
(Domning 1978) that the thin exoccipital border seen in the holotype of//, cuestae 
(UCMP 86433) is abnormal for the species. The paroccipital process is massive and 
rugose. The occipital condyle is very large and broad, and projects well abaft the occiput. 
Its upper part curves forward only slightly and there is no supracondylar fossa, again 
confirming the abnormality of the holotype. The condyle measures 78 mm in width 
and 1 17 mm front to back, a ratio of 0.67 (intermediate between values reported for 
Dusisiren and H. gigas by Domning 1978). 

Basioccipital: Completely fused with basisphenoid and exoccipitals, indicating 

Basisphenoid: Largely removed by bulldozer, but originally much thicker than in 
the (abnormal) holotype. 

Squamosal: The sigmoid ridge is about as reduced as in H. gigas, and is invisible 
in posterior view. The cranial portion dorsal to the zygomatic root is very slightly 
concave, as in the holotype. There is a protuberance about 1 cm high just above the 
external auditory meatus, an individual peculiarity not observed in the related forms. 
The temporal condyle is broad, smooth, and not distinctly demarcated; the postarticular 
fossa is broad and well developed; and the postglenoid process is about 2 cm high and 
very robust, unlike H. gigas. The processus retroversus is straight and very prominent, 
with a deep posterior indentation as in the holotype. The posterodorsal edge of the 
zygomatic process is damaged, giving the process a more lozenge-shaped outline than 
it originally had; it may have approached that of H. gigas in convexity. The forward 
end of the process is missing; its lateral edge was sharper than its medial. Its underside 
bears a clear impression for the zygomatic process of the jugal, which extended back 
almost to the level of the forward edge of the zygomatic root. Just posterior to the tip 
of the jugal is a large smooth convexity not so prominent in other Hydrodamalis 

Periotic: Present on right side but not prepared. 

Tympanic: A fragment is present on the right side, just posterior to the auditory 

Mandible. — ThQ anterior end of an immature left mandible (SDSNH 24684). badly 
worn, shows a H. gigas-\\\iQ convex outline of the anteroventral border. The mental 
foramen appears to have lain well forward (at the level of the symphysis), as assumed 
for the holotype of H. cuestae. 

Vertebrae.— Ttn of the anterior vertebrae are known from SDSNH 23719, an 
individual smaller than SDSNH 23726. Partial thoracics from other animals (SDSNH 
24679, 24680) are also available. 

Atlas (Fig. 2f-h, Table 2): The upper arch lacks a keel and articular surface for 
the axis, but has an anterior median notch flanked by a pair of rugose protuberances, 
and is penetrated posteriorly by a vertical canal about 4 mm wide. The canal for the 
first cervical nerve atop each cotyle is deep but not bridged by bone. A possible vestige 
of a vertebral arterial canal on each side, filled with matrix, may not have been patent; 
in any case there is no distinct notch on the transverse process as in Dusisiren jordani. 

Axis (Fig. 3a-c, Table 3): The odontoid process bears a broad, smooth, saddle- 


Table 2. Measurements of atlas of Hydrodamal is cuestae (SDSNH 237 1 9) from the San Diego Formation, 
in millimeters; e = estimated. 

External height 

Internal height 

Total breadth 

Width between tips of processes for transverse ligament 

Breadth across anterior cotyles 

Breadth across posterior cotyles 

Length in dorsal midline 

Length in ventral midline 








Table 3. Measurements of axis of Hydrodamalis cuestae (SDSNH 237 19) from the San Diego Formation, 
in millimeters; e = estimated. 

Total height 

Tip of odontoid process to rear of centrum 

Breadth across cotyles 

Breadth of cotyle 

Height of cotyle 

Posterior breadth of centrum 

Posterior height of centrum 

Width of neural canal 

Height of neural canal 

Breadth across postzygapophyses 

196 + 








Table 4. Measurements of cervical and thoracic vertebrae of Hydrodamalis cuestae (SDSNH 23719) from 
the San Diego Formation, in millimeters; e = estimated; a = asymmetrical. 



Tl ?T2 ?T3 

7X4 ?T5 ?T6 

Total height 

Breadth across transverse processes 
Anterior breadth of centrum 
Posterior breadth of centrum 
Height of centrum in midline 
Thickness of centrum in midline 
Width of neural canal 
Height of neural canal 
Breadth across prezygapophyses 
Breadth across postzygapophyses 
Length from front of prezygapophysis 
to rear of postzygapophysis 



219 + 



344 + 

345 + 






















































82+ 105e llOe 

shaped articular surface ventrally. The vertebral arterial canal is broadly open as in the 
holotype. The neural spine is not preserved. 

Cervicals 6-7 (Fig. 3d-e, Table 4): The vertebral arterial canal is large in C6 and 
on the right side of C7, but open laterally on the left side, which lacks the fused cervical 
rib; an anterior demifacet and a tubercular articulation, apparently for the cervical rib, 
are present. A similar asymmetry was observed in one D. Jordan! vertebra (Domning 
1978:pl. 9, fig. h). The transverse process of C6 juts farther laterad than in the corre- 
sponding vertebra of D. jordani. 

Thoracics l-?6 (Fig. 3f-g, 4, Table 4): The anterior thoracic centra have square 
sagittal sections like H. gigas and unlike the holotype, and sieve-like epiphyses as in 
the Recent species. The neural arch of Tl does not have "shoulders" like D. jordani 


Figure 3. Hydrodamalis cuestae. Vertebrae. SDSNH 23719. a-c, axis, a, anterior view; b, lateral view; c, 
posterior view; d, C6, posterior view; e, C7, posterior view, f-g, Tl. f, posterior view; g, lateral view. Scale = 
4 cm. 





- ri.r 


Figure 4. Hydrodamalis cuestae. thoracic vertebrae. SDSNH 23719. Posterior views: a, ?T2; b, ?T3; c, 
?T4; d, ?T5; e, ?T6. f, ?T6, lateral view. Scale = 4 cm. 


Figure 5. Hydrodamalis cuestae. Immature right humerus. SDSNH 24685. a, anterior view; b, medial 
view. Scale = 4 cm. 

or the holotype. The protuberances for semispinaUs tendons are indistinct. The neural 
spines are incUned backward and are much thicker posteriorly than at their anterior 
edges, in the manner of//, gigas rather than Dusisiren. The posterior sides of the spines 
are concave with distinct median ridges. The apices of the neural canals are not slit- 
like. SDSNH 24680, a partial neural arch, represents an immature animal and was 
apparently not fused to its centrum. A poorly preserved pair of thoracic vertebrae with 
arches and centra unfused (SDSNH 24679) represents a still younger animal; the max- 
imum breadth of one of these centra is 127 mm, and its thickness is 48 mm. 

Ribs. — One nearly complete right adult rib (SDSNH 24683, Fig. 7a) and several 
fragmentary ones (SDSNH 21685, 24682, 24687) have been recovered from the San 
Diego Formation. SDSNH 24683 is from near the middle of the thorax; it lacks the 
capitulum, but would have measured about 1002 mm in total straight-line length. At 


Figure 6. Hydrodamalis cuestae. Adult braincase, SDSNH 23726. a, dorsal view; b, ventral view; c, lateral 
view; d, posterior view. Left occipital condyle restored. Scale = 10 cm. 


Figure 7. Hydrodamalis cuestae. a, adult right rib. SDSNH 24683. Scale = 10 cm. b, immature distal ?right 
radius and ulna, ?medial view. SDSNH 24681. Scale in cm. 

the middle of the shaft it measures 88 x 68 mm; the distal end is swollen to 98 x 78 
mm, and slightly swept back. The distal tip bears an oval rugose concavity, and there 
is no well-marked angle. There is also no zone of cancellous bone visible on the medial 
surface. SDSNH 24682 (seven associated adult rib fragments) includes a damaged 
proximal end and two complete distal ends; the latter taper more gradually than SDSNH 
24683 and are more flattened mediolaterally (90 x 39 mm, 25 cm from distal end of 
one fragment). Except for the proximal end and another fragment of a distal tip, these 
fragments are all composed wholly of dense bone. Another fragment (SDSNH 24687) 
from the middle of a large rib (92 x 72 mm) has a large zone of cancellous bone just 
beginning to crop out on its medial side. 


Radius- ulna. — SDSNH 24681 (Fig. 7b) comprises the distal ?right radius and ulna 
of an immature animal, lacking the epiphyses. The shaft of the radius is flattened 
anteroposteriorly (about 60 x 40 mm) and is concave both mediolaterally and prox- 
imodistally on its posterior surface; it is also slightly bowed (concave ?medially). Its 
distal end is expanded to a width of 78 mm and an anteroposterior thickness of 75 
mm, with a flat posterior surface. The anterior surface bears three low, irregular knobs 
about 1.5, 4, and 10 cm, respectively, from the distal end. The shaft of the ulna is 
compressed mediolaterally and markedly bowed (concave posteriorly); its expanded 
distal end measures 71 mm mediolaterally and an estimated 75-80 mm anteroposte- 
riorly, with a flat anterior surface. The distal thickness from the anterior side of the 
radius to the posterior side of the ulna, allowing for the interosseous space whose width 
is indicated by fragments of matrix, was about 155 mm. 


The specimens herein described are distinguished from Dusisiren Domning 1978 
by the following derived characters: In the case of those from the San Mateo Formation, 
their large adult size, reduced juvenile dentition, and invisibility of the infraorbital 
foramen in ventral view; in the case of those from the San Diego Formation, their 
large size, lack of a dorsal peak to the foramen magnum, reduced sigmoid ridge on the 
squamosal, lack of a dorsal articulation between the atlas and axis, stouter and more 
posteriorly inclined thoracic neural spines, presence of a core of cancellous bone in 
immature ribs, and greater curvature of the ulna. Primitive characters of these speci- 
mens separating them from Hydrodamalis gigas include: A square cranial vault with 
more distinct temporal crests, less expanded occipital borders, a posteriorly notched 
zygomatic root, a large postglenoid process, a more anteriorly located mental foramen, 
and retention of teeth in the juvenile (at least in the Hemphillian specimens). In some 
details (zygomatic process possibly with rounded posterior end, thoracic centra square 
in sagittal section, first thoracic neural arch without "shoulders") the new specimens 
from the San Diego Formation are more derived than previously known H. cuestae, 
while in another respect (well-developed postarticular fossa of squamosal) they are 
more primitive. However, such characters may be expected to vary from more "prim- 
itive" to more "derived" conditions in a single population at any given time. In most 
respects the new specimens most closely resemble the previously described specimens 
of//, cuestae. They are, accordingly, referred to that species, with the possible exception 
of the fragmentary vertebra (SDSNH 24680) from the highest unit, which is best 
regarded as Hydrodamalis species indet. 

Conclusions reached earlier by Domning (1 978) are supported by the new material: 

1. The holotype of//, cuestae is abnormal in degree of development of the exoccipital 
border, supracondylar fossa, and basisphenoid. The skull from the San Diego For- 
mation shows the conditions expected in a form phylogenetically intermediate be- 
tween D. jordani and H. gigas. 

2. Hydrodamalis grew larger in the southern parts of its range than in the marginal 
habitat of the Commander Islands. The San Diego skull is the largest of any indi- 
vidual Hydrodamalis or other sirenian ever discovered, and several postcranial 
elements also appear to set new size records. 

Additional conclusions are permitted by the new material: 

1 . The sirenian present in the San Diego Formation is indeed //. cuestae, as would be 
predicted from the age of the unit. 

2. Hemphillian-aged juveniles of H. cuestae possessed at least four upper teeth, prob- 
ably DP2-5. 


The junior author benefited greatly from discussions with Robert M. Chandler 
(San Diego Natural History Museum) concerning West Coast Tertiary avian assem- 


The cooperation of the following companies is gratefully acknowledged in allowing 
and supporting the collection of fossils on their properties: Vance Johnson, Inc., Es- 
condido, California; Watt Industries, Rancho Santa Fe, California; Financial Scene, 
San Diego, California; and The Gersten Companies, Chula Vista, California. In addition 
the cities of Oceanside, San Diego, and Chula Vista are commended for their recognition 
of the importance of salvaging and preserving paleontological resources within their 
respective jurisdictions. 

The senior author's work on this project was supported by National Science Foun- 
dation grant #DEB 80-20265. The junior author's work was supported in part by grants 
from the Parker Foundation, the Scripps Foundation, and the J. W. Sefton Foundation. 
The additional support of Joseph and Joanne Parker is also gratefully acknowledged. 

We thank L. G. Barnes, C. A. Repenning, and an anonymous reviewer for their 
helpful comments on the manuscript. 

Literature Cited 

Barnes, L. G. 1973. Pliocene cetaceans of the San 
Diego Formation, San Diego, California, pp. 
37-42 in A. Ross and R. J. Dowlen (eds.). 
Studies on the geology and geologic hazards of 
the greater San Diego area, California. San Di- 
ego Association of Geologists. 

. 1976. Outline of eastern North Pacific 

fossil cetacean assemblages. Systematic Zool- 
ogy 25(4):32 1-343. 

, H. Howard, J. H. Hutchison, and B. J. Wel- 

ton. 1981. The vertebrate fossils of the ma- 
rine Cenozoic San Mateo Formation at Ocean- 
side, California, pp. 53-70 in P. L. Abbott and 
S. O'Dunn (eds.). Geologic investigations of 
the coastal plain, San Diego County, Califor- 
nia. San Diego Association of Geologists. 

Demere, T. A. 1983. The Neogene San Diego 
Basin: A review of the marine Pliocene San 
Diego Formation, pp. 187-195 in D. K. Larue 
and R. J. Steel (eds.). Cenozoic marine sedi- 
mentation. Pacific margin, U.S.A. Society of 
Economic Paleontologists and Mineralogists. 

Domning, D. P. 1978. Sirenian evolution in the 
North Pacific Ocean. University of California 
Publications in Geological Sciences 1 18:1-176. 

. 1982. Evolution of manatees: A specu- 
lative history. Journal of Paleontology 56(3): 

Ehlig, E. L. 1979. Miocene stratigraphy and de- 
positional environments of the San Onofre area 
and their tectonic significance, pp. 43-5 1 in 
C. J. Stuart (ed.). Miocene Lithofacies and De- 
positional Environments, Coastal Southern 
California and Northwestern Baja California. 
Society of Economic Paleontologists and Min- 

Grant, U. S., IV, and H. R. Gale. 1931. Catalogue 
of the marine Pliocene and Pleistocene Mol- 
lusca of California and adjacent regions. San 
Diego Society of Natural History Memoir 1: 

Hertlem, L. G., and U. S. Grant, IV. 1944. The 
geology and paleontology of the marine Plio- 
cene of San Diego, California, Pt. 1, Geology. 
San Diego Society of Natural History Memoir 

and . 1960. The geology and pa- 
leontology of the marine Pliocene of San Diego, 
California, Pt. 2a, Paleontology. San Diego So- 
ciety of Natural History Memoir 2a:73-133. 


1972. The geology and pa- 

leontology of the marine Pliocene of San Diego, 
California, Pt. 2b, Paleontology. San Diego So- 
ciety of Natural History Memoir 2b: 143-409. 

Howard, H. 1949. New avian records for the Plio- 
cene of California. Carnegie Institution of 
Washington Publication 584:177-199. 

. 1971. Pliocene avian remains from Baja 

California. Los Angeles County Museum Con- 
tributions to Science 217:1-17. 

. 1 976. A new species of flightless auk from 

the Miocene of California (Alcidae: Mancalli- 
nae). pp. 141-146 in S. L. Olson (ed.). Col- 
lected papers in avian paleontology honoring 
the 90th birthday of Alexander Wetmore. 
Smithsonian Contributions to Paleobiology 27. 

. 1978. Late Miocene birds from Orange 

County, California. Natural History Museum 
of Los Angeles County Contributions in Sci- 
ence 290:1-26. 

1982. Fossil birds from Tertiary marine 

beds at Oceanside, San Diego County. Cali- 
fornia, with descriptions of two new species of 
the genera Uria and Cepphus (Aves: Alcidae). 
Natural History Museum of Los Angeles 
County Contributions in Science 341:1-15. 

Ingle, J. C, Jr. 1967. Foraminiferal biofacies, 
variations and the Miocene-Pliocene bound- 
ary in southern California. Bulletins of Amer- 
ican Paleontology 52(236):2 16-394. 

Keen, A. M., and H. Bentson. 1944. Check list of 
California Tertiary marine Mollusca. Geolog- 
ical Society of America Special Paper 56:1- 

Mandel, D. J., Jr. 1973. Latest Pliocene Fora- 
minifera in the upper part of the San Diego 
Formation, California, pp. 33-36 in A. Ross 
and R. J. Dowlen (eds.). Studies on the geology 
and geologic hazards of the greater San Diego 
area, California. San Diego Association of Ge- 

Marsh, H. 1980. Age determination of the dugong 
{Dugong dugon (Muller)) in northern Australia 
and its biological implications, pp. 181-201 in 
W. F. Perrin and A. C. Myrick, Jr. (eds.). Age 
determination of toothed whales and sirenians. 
Reports of the International Whaling Com- 
mission, Special Issue No. 3. 

Miller, L. 1956. A collection of bird remains from 
the Pliocene of San Diego, California. Pro- 


ceedings of the California Academy of Science 
28(16):6 15-621. 

Moyle, W. R., Jr. 1973. Geologic map of western 
part of Camp Pendleton, southern California. 
U.S. Geological Survey Open File Map, Scale 
1:48 000. 

Repenning, C. A., and R. H. Tedford. 1977. Otar- 
ioid seals of the Neogene. U.S. Geological Sur- 
vey Professional Paper 992:1-93. 

Vedder, J. G. 1972. Review of stratigraphic names 
and megafaunal correlation of Pliocene rocks 
along the southeast margin of the Los Angeles 
basin, California, pp. 158-172 in E. H. Stine- 
meyer (ed.). Pacific Coast Miocene Biostrati- 
graphic Symposium. Society of Economic Pa- 
leontologists and Mineralogists. 

Webb, S. D., B. J. MacFadden, and J. A. Baskin. 
1981. Geology and paleontology of the Love 
Bone Bed from the late Miocene of Florida. 
American Journal of Science 281:513-544. 

Whitmore, F. C, Jr., and L. M. Gard, Jr. 1977. 
Steller's sea cow {Hydrodamalis gigas) of Late 
Pleistocene age from Amchitka, Aleutian Is- 
lands, Alaska. U.S. Geological Survey Profes- 
sional Paper 1036:1-19. 

Woodford, A. O. 1925. The San Ohofre Breccia, 
its nature and origin. University of California 
Publications in Geological Sciences 15:159- 

Woodring,W. P.,andM. N. Bramlette. 1950. Ge- 
ology and paleontology of the Santa Maria dis- 
trict, California. U.S. Geological Survey 
Professional Paper 222:1-185. 

Young, J. M., and R. W. Berry. 1981. Tertiary 
lithostratigraphic variations, Santa Margarita 
River to Aqua Hedionda Lagoon, pp. 33-51 
in P. L. Abbott and S. O'Dunn (eds.). Geologic 
investigations of the coastal plain, San Diego 
County, California. San Diego Association of 



Volume 20 Number 13 pp. 189-24Qj^/^ 20 November 1984 

Fossil Syncarida ju^ ^A/^y^^O(^ 


Frederick R. Schram 

Department of Geology. San Diego Natural History Museum, San Diego, 

Abstract. All known fossil syncaridans are reviewed, and their family level ta^oRbmy revised to 
form a more natural system. One anaspidid anaspidacean is known, Anaspidites antiquus (Chilton), 
from the Triassic of Australia. The northern hemisphere Paleozoic Palaeocaridacea are sorted into four 
families: Minicarididae (Minicaris brandi Schram, Erythrogaulos carrizoensis new genus, new species), 
Acanthotelsonidae {Acanthotelson stimpsoni Meek & Worthen, A. kentuckiensis new species, Uronectes 
fimbriatus (Jordan), U. kinniensis Schram & Schram, Palaeosyncaris dakotensis Brooks, P. micra new 
species), Palaeocarididae (Palaeocaris typus Meek & Worthen, P. retractata Caiman, P. secretanae new 
species), and Squillitidae (Squillites spinosus Scott, Praeanaspides praecursor Woodward, Nectotelson 
krejcii (Fritsch)). Several taxa are too incompletely known to be placed with certainty at this time within 
these families: Pleurocaris annulatus Caiman, Williamocalmania vandergrachti (Pruvost), Brooksyn- 
caris canadensis (Brooks), Palaeorchestia parellela (Fritsch), and Clarkecaris brasilicus (Clarke). An 
analysis of phylogenetic relationships of syncaridan families is presented. 


It is a historical curiosity that syncarids were known as fossils 45 years before they 
were discovered living in Tasmania. The understanding of their relationship to other 
eumalacostracans has unfolded only gradually, and is still not completely resolved (see 
e.g.,Dahl 1983; Hessler 1983; Schram 1981c. 1984; Watling 1981, 1983). The syncarids 
remain one of the most singularly interesting groups within the Eumalacostraca. 

The first syncarid, a Permian fossil, was described by Jordan (1 847) as Gampsonyx 
fimbriatus (now known as Uronectes fimbriatus). The species was immediately recog- 
nized by Jordan as unusual, though he compared it to amphipods in terms of its general 
form. The question of its exact systematic position, however, could not be definitively 
resolved, as evinced by Burmeister (1 855) who, in a detailed consideration of the beast, 
made passing mention of possible stomatopod a/7^amphipod similarities. Burmeister 
remarked that its closest affinities seemed to be with schizopods, yet concluded it was 
an example of a singular group ("sie ist vielmehr der Reprasentiert einer besondem 
Gruppe," p. 200). Roemer (1856) had no such reservations and placed this species 
within the Stomatopoda. 

Subsequently, Meek and Worthen (1865) described 2 more "syncarid" species, 
Acanthotelson stimpsoni and Palaeocaris typus, and placed them within the Isopoda. 

Fritsch (1870) described what he thought was a species related to G. fijnbriatus, 
which he called Gampsonychus krejcii. Fritsch (1876) also described what he thought 
was yet another species o^ ""Gampsonychus,'''' which was later placed by Zittel (1885) 
in a separate genus, Palaeorchestia parallela. 

It was Packard (1885, 1886a) who finally recognized in part the separate status of 
these fossils, and erected the taxon Syncarida. However, he placed only A. stimpsoni 
within this new group. He then proceeded to compare ""Gampsonyx'' with Palaeocaris 
typus, and concluded that these latter taxa served ". . . to bridge over the chasm existing 
between the thoracostracous suborders, Syncarida and Schizopoda . . .," (Packard 1886a: 


When Thompson (1893, 1894) described the living species Anaspides tasmaniae 
he placed it in a separate family of the Schizopoda. However, it was Caiman (1896) 
who realized the relationship oi Anaspides to the various fossil forms and united them 
altogether in the Syncarida, which he later (1904) elevated to superorder status within 
the Eumalacostraca. This arrangement completely overshadowed Grobbea's (1919) 
attempt to erect a subdivision Anomostraca within the Malacostraca for Anaspides. 

As if to celebrate this apparent resolution of syncarid affinities, a whole host of 
new fossil species soon entered the literature: Praeanaspides praecursor Woodward, 
1908; Pleurocaris annulatus Caiman, 1911; Anaspides brasilicus Clarke, 1920; Palaeo- 
caris vandergrachti Pruvost, 1922; Anaspides antiquus Chilton, 1929; Palaeocaris re- 
tractata Caiman, 1932 (actually known since 1911); and Squillites spinosus Scott, 1938 
(a name which mistakenly resurrected the idea of supposed affinities to stomatopods). 

The taxonomy of the group then achieved a certain degree of stability until Brooks 
(1962(2, b) recognized distinct generic status for Anaspidites antiquus and Clarkecaris 
brasilicus from Anaspides, and also recognized at that time the separate status of the 
Paleozoic taxa with his order Palaeocaridacea. Brooks {\962b) went on to describe a 
new species, Palaeosyncaris dakotensis, but mistakenly synonymized (Brooks 1969) 
most of the, until then separate. Palaeozoic genera with the genus Palaeocaris. 

A major revision of the fossil syncarids began with a redescription of Squillites 
spinosus by Schram and Schram (1974). Schram (1979(2) continued this review by 
reestablishing the separate generic status of several of the Paleozoic taxa, at least for 
the British Carboniferous fauna, as well as describing the earliest syncarid, Minicaris 
brandi. A second species of Uronectes, U. kiniensis, was described by Schram and 
Schram (1979). The work herein completes this revision, and examines all the known 
fossil syncarids. In addition to reestablishing as valid some old generic names, 4 new 
species are described, and 3 new genera are recognized. The artificial familial arrange- 
ment of Brooks (1962(3) is essentially discarded and a new classification of the Paleozoic 
families is put forth, one which is felt to be more natural. 


Prefixes of catalog numbers for various institutions are as follows: 

AM Museum d'Histoire Naturelle, Autun, France 

B Museum d'Histoire Naturelle, Paris, France 

BS Bayerisches Staatssamlungen fur Palaontologie und historisches Geologic, 

Munich, West Germany 

CGH Narodni Museum, Prague, Czechoslovakia 

F Australian Museum, Sydney, New South Wales 

GSE Institute of Geological Sciences, Edinburgh, Scotland 

GSL Institute of Geological Sciences, Leeds, England 

I, In British Museum (Natural History), London, England 

ISGS Illinois State Geological Survey, Urbana, Illinois 

Jk Museum fur Naturkunde (Janensch Catalog), Berlin, East Germany 

M, Me Narodni Museum, Prague, Czechoslovakia 

NB Rijks Geololgische Dienst, Heerlen, The Netherlands 

NYSM New York State Museum, Albany, New York 

PE Field Museum of Natural History, Chicago, Illinois 

PMB Museum fiir Naturkunde (Paleontologisches Museum Catalog), Berlin, East 


SDSNH San Diego Natural History Museum, San Diego, California 

US University of Sydney, Paleontology Collection, Sydney, New South Wales 

USNM National Museum of Natural History, Smithsonian Institution, Washington, 


X University of Illinois, Paleontology Collection, Urbana, Illinois 

YPM Yale Peabody Museum of Natural History, New Haven, Connecticut 


Living Anaspidacea 

In 1980 I was able to collect and study several species of living anaspidaceans in 
Tasmania with the assistance of the staff of the University of Tasmania. Several of 
these observations have not been recorded before, and offer some insights into the 
biology of the fossil syncarids. 

The most widely dispersed anaspidacean is Anaspides tasmaniae (a rather variable 
taxon which will probably contain several subspecies— R. Swain, personal communi- 
cation) which occur in streams, lakes, and caves widely scattered about the island of 
Tasmania. The animals are in constant motion, somewhat less so in the wild than in 
the laboratory. They tend to engage in a constant and random patrol of their pools. 
They seem to ignore each other, and in the laboratory they climb over each other in 
the course of their wanderings like any other obstacle in their path. The exopods 
constantly vibrate anteriorly to posteriorly, moving the epipodites in the process. The 
annulate pleopods are directed ventro-laterally, and push the body along the bottom 
in metachronal rhythm with the thoracic endopods. While collecting Anaspides, some 
inadvertently fell out of the dip net onto the ground, whereupon they righted themselves 
and commenced to explore their terrestrial environs with ease. Richardson {personal 
communication) relates that occasionally, albeit rarely, they are naturally encountered 
out of their pools on land. This species does not swim at all well. When startled they 
will execute a single caridoid flexure that propels them up into the water column. 
However, they then drift passively until gravity returns them to the bottom. This 
probably accounts for their inability to survive in areas where European sport fish have 
been introduced, and also makes it extremely easy to collect them. Overall, A. tasmaniae 
is a very alert animal. The flagellae of both sets of antennae orient in different directions 
and constantly sweep about. They are omnivorous, prefering to scavenge, and are also 
known to pick up large sand grains and manipulate them with their mouthparts, ap- 
parently to scrape them of organics. 

Paranaspides lacustris is a smaller animal than A. tasmaniae, and exhibits some 
distinctly different behavior. Their pleopods also function like those o{ Anaspides when 
they walk on the bottom. However, Paranaspides seems to be much more versatile in 
its locomotion. When startled they may execute the single caridoid flexure already 
mentioned, or they may dart away in some direction parallel to the bottom, or they 
may lay quite still. When they do enter the water column they are capable of swimming 
quite well. Paranaspides was observed by me to swim for hours near the surface in a 
small thermos-container. In the laboratory, they were generally less active animals than 
Anaspides. Although they have a flexure point in the abdomen to facilitate the caridoid 
reaction, when they rest in their habitat on the bottom ooze or on the water plants 
they prefer, they reflex and hold the tailfan dorsad off the substrate. 

Allanaspides is the smallest of the Anaspididae. A. helionomus is collected from 
yabbie (crayfish) burrows. Unlike the 2 species above, A. helionomus does not beat its 
exopods in a simple to-and-fro pattern, but seems to rotate them. The exact manner 
is not clear, but the resultant current sucks water under the head of the animal and 
back towards the tail. Small, young individuals do not beat their exopods when at rest; 
only the adult animals do. The anterior thoracopods are oriented anteriorly under the 
head while at rest; in combination with the described current they may function in 
filter feeding. Also, unlike Anaspides and Paranaspides, Allanaspides helionomus uses 
its pleopods in a somewhat different manner. The uniramous annulate abdominal 
appendages are held rigid and each pair is oriented in a different direction. When they 
assist in walking they push off the bottom like oars using just their tips. Walking 
in A. helionomus is best described as a "scurrying crawl," occurring in intermittent 
bursts of activity. When at rest, the first 2 pleopods vibrate vigorously to aerate the 
fleshy thoracic epipodites. Allanaspides swims very well, and was observed to do so 
even upside down. The animals may be detritovores since in the laboratory they were 
observed to fondle fecal pellets with their mouthparts. The burrows these animals live 
in occur in grass marshes on surfaces of gentle slope, aflTording a modicum of drainage 
and no long-standing water. 


Micraspides calmani possess a very flexible body, easily achieving flexion dorsally 
and ventrally as well as considerable lateral bend, and is the most infaunal of any of 
the species observed by me in Tasmania. It also lives in pools and yabbie burrows in 
grass swamps, but seems to better tolerate conditions with poorer drainage than do 
species of Allanaspides. Micraspides moves with bursts of intermittent scurrVing, rem- 
iniscent of that seen in some centipedes. The annulate pleopods are held somewhat 
stiffly, and operate only within the metachronal sequence of all the limbs, thus differing 
only in form and not function from the thoracopods. When not moving they do not 
move any of the appendage parts. No caridoid escape reaction could be elicited from 
Micraspides; when startled or prodded the animals would take evasive action by turning 
laterally or flexing ventrally, eventually to change their direction of movement 1 80°. 
They are thus ideally adapted to climbing in, around, over, under, and through obstacles 
in the vegetation-choked, muddy habitats they prefer. 

Several aspects of the above have direct bearing on interpretation of the fossils. 
The annulate pleopods seen on the living forms are noted as one of the most versatile 
and important aspects of the anatomy of these creatures; serving to achieve walking, 
swimming, and ventilation of epipodites. They also form, for the most part, a functional 
continuum with the thoracopods (MacMillan et al. 1981). This has great bearing on 
the Paleozoic fossils, which were once mistakenly thought (Brooks 1962/?) to all have 
flap-like pleopods and thoracic exopods. The functional system for pleopods seen in 
the living forms is quite distinct from what might be postulated for those few paleo- 
caridaceans with biramous flap-like pleopods, which would appear to have been capable 
of only one action, a to-and-fro vibration on the ventral side of the abdomen. Such 
limbs would serve in swimming, but have little or no effect on walking on the bottom 
or producing ventilatory currents over the thoracic epipodites. The latter would have 
to be achieved by the vibration of the flap-like thoracic exopods. In turn, the exopods, 
because of their form and consequent limitation of movement, could not serve to 
generate potential filtering currents around the body as do the rotatory exopod move- 
ments of a form like Allanaspides. 

Palaeocaridaceans such as the acanthotelsonids, or in part the palaeocaridids, 
probably exhibited a functional system not unlike that seen in other eumalacostracan 
groups, such as mysidaceans, euphausiaceans, and natant decapods, wherein the pleo- 
pods are the sole or primary organs of swimming in the adult stage. The system seen 
in the anaspidaceans, the squillitids, and in part the palaeocaridids would then appear 
to possibly represent a functional advance in which the entire trunk appendage series 
is capable of acting as a coordinated unit. In this respect, it seems to have been a 
successful enough arrangement of parts to have perhaps evolved at least twice within 
the syncarids: once in the palaeocaridid/squillitid line and again in the anaspidaceans. 

Some interesting questions arise for which, at the present, there are no obvious 
answers. In those living eumalacostracans for which the use of pleopods for swimming 
is well developed there has evolved an excellent caridoid escape reaction. Does the 
existence of a similar anatomical system in some of the Palaeocaridacea mean that 
they too may have had a well-developed caridoid escape reaction, in contrast to the 
rather inefficient single-flexure behavior seen in living anaspidaceans? Dahl (1983) 
suggests that the caridoid escape reaction of eumalacostracans was independently evolved 
in mysidaceans and natant eucarids. Does its possible existence in some palaeocari- 
daceans mean there was a third independent evolution of this behavior, or is its possible 
presence in the syncarids an argument for the caridoid escape reaction being considered 
as a derived character applicable to all eumalacostracans (Hessler 1983) and which has 
merely been repeatedly lost? Do these functional considerations tell us anything about 
character polarities (see next section) within syncarids? If the integrated system with 
annulate appendage parts represents a functional advance within the syncarid line, this 
might indicate that the purely flap-like structures are primitive and that animals which 
possess them are closer to the stem-group. Thus bathynellaceans and many of the 
palaeocaridaceans might be considered more primitive than anaspidaceans. On the 
other hand, if annulate limb parts and a functionally integrated trunk might represent 










Figure 1 . Classical arrangement of the superorders of Eumalacostraca sensu stricto. Derived characters are: 
1) caridoid escape reaction (and its associated features of abdominal specializations, see Hessler 1983), 2) 
antennal scale of a single joint, 3) loss of a carapace, 4) carapace fused to thoracomeres, and 5) oostegite 
brood pouch. 

an advance for syncarids, another scenario is possible. There is a tendency for pae- 
domorphosis in syncarids (reduced or absent posterior limbs, small body size, free first 
thoracomere). Flap-like limb parts and a restriction of the pleopods to a swimming 
behavior may represent a retention of "larval" features, and thus provide further 
evidence for structural and behavioral paedomorphosis with palaeocaridaceans and 
bathynellaceans being the more derived groups. 

Higher Taxonomy And Phylogeny 

Problems arise in attempting to assess relationships of taxa within the syncarids. 
However, these are no more difficult than the problems associated with attempting to 
assess the position of syncarids in relation to other eumalacostracans. What are the 
unique characters which define a taxon Syncarida? In the classic scheme of Caiman 
(Fig. 1) Eumalacostraca sensu stricto are principally characterized by their caridoid 
escape reaction (1) and 1 -jointed antennal scale (2). The syncarids are a sister group of 
peracarids and eucarids, defined by a derived feature (3), loss of the carapace (a condition 
paralleled by a similar loss in the line leading to amphipods and isopods). No opposing 
shared derived characters, however, join eucarids. with their carapace fused to the 
thoracomeres (4), and the peracarids, with their oostegite brood pouch (5). 

Schram (1981) and Watling (1981, 1983) have taken up the problems engendered 
by the Caiman system and have offered differing solutions to those difficulties. The 
Watling model has difficulties in providing shared derived characters at the higher 
taxonomic levels, and won't be dealt with further here. The system proposed by Schram 
(1981) had the syncarids as a sister group to the isopods and amphipods. A difficulty 
with that scheme is that it left no derived features to define the syncarids. A subsequent 
cladistic analysis (Schram, in press) utilizing 31 characters and a Wagner 78 program. 


4artices in 
thoracic endopods 

5 articles in 
thoracic endopods 

first thoracomere 


Pa aeocaridacea 

first thoracomere 
fused to cephalon 


Figure 2. Baupldne one can recognize with the Syncarida. The combination of the first thoracomere fused 
to the cephalon with 4-segmented thoracopodal endopods was apparently never realized. 

while confirming taxa based on Baupldne derived from consideration of only 3 char- 
acters (Schram 1981), does not second the linking of syncarids with amphipods and 
isopods. This more recent analysis, however, does generally indicate that Syncarida is 
a Gilmour-natural taxon, and also reveals that syncarids probably are very primitive 

Indeed, several aspects of the biology of living forms would reinforce this conclu- 
sion. Although the living anaspidaceans have a caridoid escape reaction, it is imperfectly 
developed. It typically consists of a single flick of the abdomen that projects the animal 
up into the water column, after which the animal either passively floats or slowly swims 
back to the bottom. This is in contrast to the strong caridoid escape reaction seen in 
the mysidaceans and eucarids. In addition, syncarids lay their eggs free and gastrulate 
by involution into a blastocoel— both primitive features. However, the development 
within the egg proceeds to hatching at a rather advanced, free-living stage (early zoeal 
larval type in bathynellaceans, or miniature version of the adult in anaspids), which 
are generally considered derived features. 

Syncarids are thus generally considered to be a monophyletic taxon. However, 
discerning possible phylogenetic relationships within the group poses problems. One 
could recognize 3 basic morphotypes or Baupldne (Fig. 2) within the syncarids: a free 
first thoracomere and less than 5 segments in the thoracopodal endopods (Bathynel- 
lacea), a free first thoracomere and 5-segment thoracic endopods (Palaeocaridacea), 
and a first thoracomere fused to the cephalon and 5 segments in the thoracic endopods 
(Anaspidacea). The characters used here are the "traditional" ones used for decades to 
sort major syncarid groups. 

Problems arise when one is forced to choose between delineating syncarids pri- 
marily on the basis of endopodal segment numbers or on the basis of degree of fusion 
of the first thoracomere into the cephalon. Different cladograms and classifications of 
the syncarids also result based on whether palaeocaridaceans are to be perceived as a 
monophyletic or paraphyletic group. 

The traditional approach (Fig. 3) (Brooks 1969, Schminke 1975) essentially dis- 
criminates between the bathynellaceans with 4-segmented thoracic endopods (1) and 
palaeocaridaceans and anaspidaceans which have 5-segmented thoracic endopods. In 
addition, bathynellaceans have the incisor process of the mandible fused to the tooth 
row (2), and the eighth thoracopod of males modified for copulation (3). Character (2) 
is difficult to assess in the palaeocaridaceans, since the mandibles generally are not 
preserved well enough to be able to evaluate whether they are truly primitive in form 
(the well-preserved massive mandibles seen in Palaeocaris, with its distinct incisor 
process, may or may not be indicative of all palaeocaridaceans). The palaeocaridacean- 


Figure 3. A "traditional" presentation of relationships within Syncarida. Derived characters are: 1 ) 4-segment 
thoracic endopods, 2) incisor process fused to tooth row, 3) eighth thoracopod copulatory, 4) furcae lacking, 
5) first thoracopod typically modified, 6) "precoxae" lacking, 7) paragnaths lacking, 8) diagonal spine row 
on uropodal protopod, 9) first thoracopod reduced, 10) first thoracomere fused to cephalon, 11) eighth 
thoracopod not parallel to anterior thoracopods, 1 2) greatly shortened maxillipedal palp, 1 3) first thoracomere 
generally reduced in size, 14) reduction and/or lack of pleopods, 15) pleotelson, 16) anterior thoracopods 
rapacious, 17) annulate pleopods, 18) massive mandibles, 19) annulate thoracic exopods, 20) large, setose 
uropod protopod, 21) rostrum separated, 22) maxillule palp lacking, 23) maxillulary palp hook-like. 24) 
mandibular incisor process lacking, 25) endite lobe on first thoracopod, 26) maxilla proximal endite lacking 
spine, 27) anus terminal, 28) mandibular palp lacking. 

anaspidacean line is delineated by several synapomorphies: lack of caudal furcae (4), 
a first thoracopod typically modified in some manner (5), and lack of thoracopodal 
precoxae (6). This last character is difficult to assess, since whether the presence of this 
feature in some bathynellaceans is really a true precoxal leg joint or just an articulating 
ring on the body is not clear. 

Within the bathynellaceans the Parabathynellidae are characterized by the lack of 
paragnaths (7), while the Bathynellidae are marked by a uropodal protopod with a 
diagonal row of spines (8). 


The palaeocaridaceans are characterized by a derived feature that is difficuU to 
evaluate. Schminke (1975) was the first to point out that the first thoracopod is ap- 
parently reduced in size as well as number of joints in the endopod (9). While this is 
clearly true for palaeocarids, acanthotelsonids and some squillitids, the form in mini- 
carids is incompletely known. In the latter family the first thoracopod is large, but it 
is unclear if there is a complete array of 5 segments in the endopod. However, in the 
squillitid genus Nectotelson there is some evidence that indicates the carpus of the first 
thoracopod is apparently not as large as on thoracopods 2 through 8. Generally, in the 
former 3 families, besides the greatly shortened maxillipedal palp (12) the first thorac- 
omere is reduced to some degree (13). 

Minicarididae are very small animals which appear to have a reduced number of 
pleopods (14). Admittedly this observation could be due to vagaries of preservation; 
however, the rest of the body and appendages of these creatures are preserved well 
enough to discern all pertinent features of their structure. The possible presence of a 
pleotelson among these genera ( 1 5) might eventually serve to further define this group. 

The Acanthotelsonidae are clearly delineated by the specialized, rapacious form 
of their anterior thoracopods (16). Within that family, Uronectes has only the second 
thoracic appendages so modified, while Acanthotelson and Palaeosyncaris have the 
second and third so specialized. The styloid telson and uropods delineate the species 
of Acanthotelson, whereas a robustly spinescent telson and lateral margin of the uro- 
podal exopod characterize Palaeosyncaris. 

The palaeocarids and squillitids are characterized by annulate pleopods (17). The 
Palaeocarididae, though for the most part rather generalized, appear to be distinguished 
by rather massive mandibles ( 1 8). The Squillitidae are unified by their acquisition of 
annulate thoracic exopods (19). Squillites has uniramous pleopods, Nectotelson and 
Praeanaspides have biramous pleopods, and Praenaspides has a distinctive rectangular 
and laterally spinose telson. 

Schminke (1975) has clearly analyzed the distribution of characters within the 
Anaspidacea, and these are repeated here only for completeness. Anaspidaceans are 
united in possessing a first thoracomere completely fused to the cephalon (10) and the 
eighth thoracopod offset at an angle and not parallel to the other thoracopods (11). The 
anaspids and koonungids have a large and setose uropodal protopod (20), while psam- 
maspids and stygocarids have a separated rostrum (21) and lack a palp on the maxillules 
(22). Anaspididae are distinguished by having the maxillulary palp reduced to a hook- 
like spine (23), and the Koonungidae lack an incisor process on the mandible (24) and 
have an endite lobe on the first thoracopod (25). The Psammaspididae have no spine 
on the proximal endite of the maxillae (26), while the Stygocarididae have a terminal 
anus (27) and lack a mandibular palp (28). 

A classification of Syncarida produced from the above analysis is similar to that 
usually encountered for the group, except that a more natural array of palaeocaridacean 
families is established than that used by Brooks (1962a, b, 1969). 

Order Syncarida Packard, 1885 

Suborder Bathynellacea Chappuis, 1915 

Family Bathynellidae Chappuis, 1915 

Family Parabathynellidae Noodt, 1965 
Suborder Palaeocaridacea Brooks, 1962 

Family Minicarididae, nov. 

Family Palaeocarididae Meek & Worthen, 1865 

Family Squillitidae Schram & Schram, 1974 

Family Acanthotelsonidae Meek & Worthen, 1865 
Suborder Anaspidacea Caiman, 1904 

Family Anaspididae Thompson, 1894 

Family Koonungidae Sayce, 1908 

Family Psammaspididae Schminke, 1974 

Family Stygocarididae Noodt, 1963 


Figure 4. An alternative presentation of relationships within Syncarida. Derived characters are: 1) first 
thoracomere reduced or fused to cephalon. 2) incisor process fused to tooth row, 3) furcae lacking, 4) first 
thoracopod modified, 5) reduced and/or absent pleopods, 6) annulate thoracic exopods and pleopods, 7) 
4-segmented thoracic endopods, 8) reduced and/or absent pleopods, 9) eighth thoracopod copulatory, 10) 
diagonal spine row on uropodal protopod, 1 1) paragnaths lacking, 12) first thoracomere fused to cephalon, 
13) eighth thoracopod not parallel to anterior thoracopods, 14) greatly reduced maxillipedal palp, 15) large, 
setose uropodal protopod, 16) rostrum separate, 17) maxillulary palp lacking, 18) maxillulary palp hook- 
like, 19) mandibular incisor process lacking, 20) endite lobe on first thoracopod, 21) maxilla proximal endite 
lacking spines, 22) anus terminal, 23) mandibular palp lacking, 24) anterior thoracopods rapacious, 25) 
annulate pleopods, 26) massive mandibles, 27) annulate thoracic exopods. 

An alternative analysis of cladistic relationships (Fig. 4) can be performed for 
syncarids with initial assumptions somewhat different from those of the traditional 
system presented above. Rather than make the first dichotomy one based essentially 
on numbers of segments in thoracic endopods, one could distinguish between syncarids 
with no modification of the first thoracomere and those with a first thoracomere mod- 
ified in some way (1). The former line includes the bathynellaceans and might be further 
characterized by mandibles (2) with an incisor process fused to the tooth row (as noted 
above, a character impossible to verify as yet on all the fossils). This line divides into 
a branch leading to some fossil families which lack furcae (3) and which may have a 
first thoracopod modified from the form seen in the second through eighth thoracopods 
(4). Within this branch, the Minicaridadae apparently lack or have a reduced number 
of pleopods (5) while the Squillitidae (in part, including Nectotelson and Squillites) 
have annulate thoracic exopods and pleopods (6). 

The branch leading to the bathynellaceans of course delineates the 2 families on 
the basis of the bathynellid's possession of a diagonal row of spines on the uropodal 
protopod (10) and by the parabathynellid's lack of a paragnath (11). 

The Anaspidacea fuse the first thoracomere into the cephalon (12) and have the 
eighth thoracopod offset from the seventh thoracopod (13). The opposing branch with 
its reduced but free thoracomere and parallel seventh and eighth thoracopods possess 


a derived reduction in the first thoracopodal endopod being very short (14). The an- 
aspid/koonungid line, as above, have large and setose uropodal protopods (15); and 
within that the Anaspididae have a hook-like maxillulary palp (18), while Koonungidae 
lack a mandibular incisor process (19) and have gnathobasic lobes on the first thora- 
copod (20). The psammaspid/stygocarid line has a separate rostrum (16) and lacks a 
maxillulary palp (17). The Psammaspididae lack spines on the proximal endites of the 
maxillae (21), while the Stygocarididae have a terminal anus (22) and lack a mandibular 
palp (23). 

The Acanthotelsonidae have rapacious anterior thoracopods (24), while the pa- 
laeocarid/squillitid group has annulate pleopods (25). The Palaeocarididae have mas- 
sive mandibles (26), and Praeanaspides (a squillitid) has annulate thoracic exopods 

This analysis could yield a classification somewhat different than the traditional, 
in that essentially 4 groups can be recognized. Brooks' order Palaeocaridacea emerges 
as a polyphyletic taxon, thus the major groups might best be recognized as superfamilies. 

Order Syncarida 

Superfamily Bathynelloidea 

Family Bathynellidae 

Family Parabathynellidae 
Superfamily Minicaridoidea 

Family Minicarididae 

Family Squillitidae (in part) 
Superfamily Palaeocaridoidea 

Family Palaeocarididae 

Family Acanthotelsonidae 

? Praeanaspides 
Superfamily Anaspidoidea 

Family Anaspididae 

Family Koonungidae 

Family Psammaspidae 

Family Stygocarididae 

In many respects, this second arrangement is an unsatisfactory system. The dif- 
ferences encapsulated in these 2 classifications of the syncarids arises from a dichotomy 
involved in outgroup comparison of the "palaeocaridaceans," and in both schemes it 
involves establishing derived characters in the Paleozoic families. The problem could 
be expressed as a simple dilemma derived from initial consideration of the syncarid 
morphotypes: which is more important, the fusion of first thoracomere into the ceph- 
alon, or the loss of a joint in the thoracopodal endopods? The initial "weighting" 
determines the course of the subsequent analysis. 

In the traditional scheme (Fig. 3) 28 apomorphies are used to define the 10 families 
of Syncarida; in the alternative scheme (Fig. 4) there are 27 apomorphies to separate 
1 1 "family" level taxa. The traditional scheme thus seems to possess slightly more 
information value. It also more clearly justifies its initial dichotomy (based on joint 
number in thoracic endopods) with the greatest number of congruent features. For these 
last reasons, as well as the fact that the alternative scheme requires too many uncertain 
judgments at this time involving the poorly known minicaridoids, I have opted in the 
systematic section of this monograph to retain the traditional classification of syncarids 
into suborders. However, more detailed and exacting knowledge of the minicarids and 
Nectotelson someday may allow a more reasoned selection to be made between these 
two systems. 

Nevertheless, the problem of analyzing syncarid phylogeny is not so easily disposed 
of (as if the above taxonomic dilemma were easy). Both of the schemes above take for 
granted essentially the same position in regard to polarity of a basic character in the 
group, viz, that the primitive condition is one in which the first thoracomere is free 


and large, and that increasing specialization is achieved as this segment is reduced and 
eventually fused into the cephalon. Is this necessarily the case? 

Schminke (1981) presents a well-documented series of arguments for the progenetic 
paedomorphic derivation of bathynellaceans from some ancestral syncarid condition 
in which an adult animal, presumably of a palaeocarid or anaspid form, had a long 
larval sequence. If we extend Schminke's arguments, might we not question whether 
the anatomical stages seen in bathynellaceans (large and free first thoracomere, flap- 
like thoracic exopods, reduced number of thoracic endopodal segments, reduced or 
missing pleopods, and caudal furcae) are really primitive? Rather, might we not consider 
these features to be actually derived by the agency of progenesis from some ancestral 
adult in which none of these "larval" features were expressed. In such an interpretation, 
the most "primitive" adult state would be one in which the first thoracomere is fused 
to the cephalon, and that the manifestations of successive degrees of freedom of the 
first thoracomere are increasingly derived. 

This assumption involving a reversed polarity would lead in turn to a rather 
controversial cladistic analysis (Fig. 5). A bathynellacean/palaeocaridacean line would 
be characterized by the presence of a free first thoracomere (1), the anaspidacean line 
by the eighth thoracopod being offset from and not parallel to the seventh (2). 

The characterizations within the Anaspidacea follow those already given above: 
large, setose uropodal protopods in anaspids and koonungids (3), a separate rostrum 
(4) and no maxillulary palp (5) in psammaspids and stygocarids, a hook-like palp on 
the maxillule (6) in anaspids, lack of a mandibular incisor process (7) and endite lobes 
on the first thoracomere (8) in koonungids, lack of spines on the proximal endite of 
the maxilla (9) in psammaspids, and a terminal anus (10) and lack of a mandibular 
palp (11) in the stygocarids. 

The Bathynellacea share several advanced characters: a 4-segment thoracic en- 
dopod (12), fusion of the incisor process to the tooth row in the mandible (13), a 
copulatory eighth thoracopod in the male (14), and a first thoracomere as large as any 
succeeding thoracomere (15). The Palaeocaridacea lack furcae (16). 

The bathynellids have a diagonal row of spines on the uropodal protopod (17), 
while the parabathynellids lack paragnaths (18). 

Within the palaeocaridaceans, the minicarid/squillitid (in part) line convergently 
develops the enlarged first thoracomere (19), while the other families have a reduced 
endopod on the first thoracopod (20). The minicarids apparently have reduced or absent 
pleopods (21), while the squillitids have annulate thoracic exopods and annulate pleo- 
pods. The acanthotelsonids have rapacious anterior thoracopods (23), while the pa- 
laeocand/ Praeanaspides line has annulate pleopods (24). The palaeocarids have mas- 
sive mandibles (25) and the Praeanaspides also possess annulate thoracic exopods (26). 

The above scheme in comparision with the traditional and alternative schemes 
discussed earlier unfortunately uses only 26 apomorphies to define its end points and 
has a rather high number of convergent characters. Note, however, that the taxonomy 
which results from this cladogram is similar to that of the traditional classification, 
except that the Anaspidacea in this latter scheme are felt to be closest to the primitive 
condition, and the Squillitidae sensu stricto are separated from the genus Praeanaspides. 

Still another analysis alternative to the above is possible (Fig. 6) also involving 
the reversed polarity, but utilizing only 25 apomorphies. The anaspidoid line is as 
above. The bathynelloid/minicaridoid line has an enlarged first thoracomere ( 1 2), while 
the palaeocaridoid line has a reduced endopod on the first thoracopod. The bathynelloid 
hne has the 4-segment endopod (14), incisor process fusion (15), and copulatory eighth 
thoracopod (16) noted before; and the minicaridoid line has a problematic apomorphy 
difficult to assess because of preservation, i.e., the first thoracopod large but possibly 
not structurally identical to the second and following thoracopods (17). The family 
apomorphies [Bathynellidae (18), Parabathynellidae (19), Minicaridiae (20), Squilliti- 
dae (in part) (21), Acanthotelsonidae (22), Palaeocarididae (23. 24), and Praeanaspides 
(23, 25)] are all those noted in the schemes already discussed, especially that in Figure 5. 




^ / ■/ ^ / v^ 

/ / # / / -^ 

Figure 5. Relationships within Syncarida involving reversed polarity with loss of fusion of the first tho- 
racomere with the cephalon due to paedomorphosis. Derived characters are: 1) first thoracomere free, 2) 
eighth thoracomere not parallel to anterior thoracopods, 3) large, setose uropodal protopod, 4) rostrum 
separate, 5) maxillulary palp lacking, 6) maxillulary palp hook-like, 7) mandibular incisor process lacking, 
8) endite lobe on first thoracomere, 9) maxillary proximal endite lacking spines, 10) anus terminal, 11) 
mandibular palp lacking, 12) 4-segment thoracic endopod, 13) incisor process fused to tooth row, 14) eighth 
thoracopod copulatory, 15) first thoracomere subequal to any succeeding thoracomeres, 16) furcae lacking, 
1 7) diagonal spine row on uropod protopod, 1 8) paragnaths lacking, 1 9) first thoracomere subequal to any 
succeeding thoracomeres, 20) greatly reduced maxillipedal palp, 21) reduced and/or absent pleopods, 22) 
annulate thoracic exopods and pleopods, 23) anterior thoracopods rapacious, 24) annulate pleopods, 25) 
massive mandibles, 26) annulate thoracic exopods. 

This last cladogram coiresponds to the alternative classification given above, except 
that the superfamily Anaspidoidea is now presented as the closest to a primitive con- 
dition for Syncarida as a whole. 

What can we conclude about syncarid evolution? We should be cognizant of some 
level of uncertainty as to just how these taxa are related to each other. However, a 
more definitive resolution of the problem must await better and more detailed infor- 
mation about the Paleozoic syncarids. Characters which delineate the living and fossil 
families are not equivalent in the sense that the living families are separated on the 
basis of details of mouthparts, whereas the fossil families are largely resolved on the 
basis of gross form of trunk appendages. Ideally, more mouthpart data for Palaeocar- 
ideacea could have allowed a more complete data matrix than that used here to be 
analyzed with a Wagner 78 program. The resultant rigor could have mathematically 
determined parsimony and homoplasy. However, phylogenetic trees and taxonomies 
are pragmatic instruments (Charig 1982, Schram 1983), and the lack of any data that 
we would like to have should not be an excuse for not attempting to organize that 


Figure 6. An alternative presentation of relationships within Syncarida involving reversed polarity with 
loss of fusion of the first thoracomere with the cephalon due to paedomorphosis. Derived characters are: 1 ) 
first thoracomere free, 2) eighth thoracomere not parallel to anterior thoracopods, 3) large, setose uropodal 
protopod, 4) rostrum separate, 5) maxillulary palp lacking, 6) maxillulary palp hook-like, 7) mandibular 
incisor process lacking, 8) endite lobe on first thoracomere, 9) maxillary proximal endite lacking spines, 10) 
anus terminal, 1 1) mandibular palp lacking, 12) first thoracomere, 13) reduced endopod on first thoracopod, 
14) 4-segment thoracic endopod, 15) incisor process fused to tooth row, 16) eighth thoracopod copulatory, 
17) ? form of first thoracopod, 18) diagonal spine row on uropod protopod, 19) paragnaths lacking, 20) 
reduced and/or absent pleopods, 21) annulate thoracic exopods and pleopods, 22) anterior thoracopods 
rapacious, 23) annulate pleopods, 24) massive mandibles, 25) annulate thoracic exopods. 

information which we do have. We should simply recognize the limits of the information 
at hand, and be aware of its effect on the level of uncertainty engendered in our present 
understanding of syncarid evolution. Nevertheless, I would hope that the organization 
of the fossil Syncarida used here is more adequate than anything that we have had 


The system of annotated synonymy, summarized by Matthews (1973), is used in 
this section of the monograph. This should facilitate use and evaluation of my systematic 
decisions by any future workers. 

Order SYNCARIDA Packard, 1885 

Suborder PALAEOCARIDACEA Brooks, 1962 


Diagnosis. — Thoracic exopods unisegmental. Pleopods unisegmental, if present. 
First thoracomere large, not reduced nor fused to cephalon. 

Type genus.— Minicaris Schram, 1979. 

Remarks. — The distinctive nature of the first thoracomere and the unisegmental 
or flap-like form of the pleopods when present clearly warrants separate family status. 


Figure 7. A) Reconstruction of Minicahs brandi. scale 2 mm (redrawn from Schram 1979a); B) tailfan 
to same scale as body; C) dorsal view of right antenna slightly enlarged. 

In addition, the small size, possible absence of posterior pleopods, the possible presence 
of a pleotelson, and the early age (Lower Carboniferous) is of interest with regard to a 
parallelism to, or a possible origin of, the Bathynellacea {see Higher Taxonomy and 
Phylogeny section). 

Genus MINICARIS Schram, 1979a 

Diagnosis. — VtdunclQS of antennules and antennae subequal. At least first pleopod 
present and well developed. Uropods narrow and blade-like. (?)Pleotelson. 
Type species.— Minicaris brandi Schram, \919a. 

Minicaris brandi Schram, \919a 
Fig. 7 

v.* 1979a Minicaris brandi Schram, p. 109, figs. 52 & 53. 
1979b Minicaris brandi Schram. Schram, p. 170, table 2. 

1981 Minicaris brandi Schram. Schram, p. 131, table 2, fig. 6D. 
7952 Minicaris brandi Schram. Wood, p. 577. 

1982 Minicaris brandi Schram. Schram, p. 122, fig. 8A. 

Diagnosis. —Since there is but one species, the diagnosis is the same as that of the 

Holotype. —GSE 13056. Long Livingston Borehole no. 25, West Lothian, Scotland. 
1071-1151 foot section, below Pumpherstone Shell Bed, Lower Oil Shale Group, Di- 
nantian, Lower Carboniferous. 

Other locality.— Questionably reported from along Manse Bum, Bearsden, near 
Glasgow, Scotland, in shales equivalent to the Top Hosie Limestone, lowermost Na- 
murian (Wood 1982). 

i)€'5'cn/)//oA2.— Antennular peduncle 3 joints, proximal-most joint one-half total 
length of peduncle, distal 2 joints progressively shorter. Antennal protopod distal joint 
twice the proximal, scaphocerite oval with distal tip pointed and setose, proximal 2 
flagellar joints peduncular. Thoracomeres with rounded pleura, posterior comers acute. 
Thoracopodal exopods narrow. All thoracopods appear equal, ischium long, merus and 
carpus short, propodus moderate, dactylus short. Abdominal pleura rounded. If not a 
tme pleotelson, telson not sharply sutured from sixth pleomere. Telson spade-like, 
setose. Uropods blade-like, setose, possibly with diaeresis. 

/^emar/:^. — Reexamination in 1980 of the holotype, and still only good specimen 
of this species, confirmed all the pertinent points of the anatomy above. The lack of 


Figure 8. A) Reconstruction of Erythrogaulos carrizoensis, scale 2 mm; B) tailfan to same scale as body. 

all but the first pleopod in the abdominal series is still not completely understood. It 
is possible the more posterior pleopods were not preserved, but it is also possible that 
they were never there to begin with. The small size (8 mm) and general form of the 
animal might indicate a possibility of paedomorphosis in the evolution of this taxon, 
since so many of the living small syncarids do not develop complete series of pleopods 
nor completely separate the telson from the last pleomere. Only more and better material 
can allow us to choose between these alternatives. 

Genus ERYTHROGAULOS new genus 

Diagnosis. — 'PosXtnov comers of pleomere pleura serrate. Telson distally spinose. 
Uropodal exopod distally spinose. 

Type species.— Erythrogaulos carrizoensis new species. 

Etymology.— A. reference to the stratigraphic horizon, Red Tanks Member, Madera 
Formation, Lower Permian. 

Erythrogaulos carrizoensis new species 
Fig. 8; Plate 1, figs. A & B 

Diagnosis. SincQ there is but one species, the diagnosis is the same as that of the 

//o/o/yp^.-SDSNH 25141 (Plate 1, figs. A «fe B). Carrizo Arroyo, Lucero Mts., 
southeastern Valencia County, New Mexico. Upper Red Tanks Member, Madera For- 
mation, Wolfcampian, Lower Permian. (Collected by Dr. Jarmilla Kukalova-Peck, 
Carlton University, Ottawa, Ontario, Canada.) 

Etymology. — MXqt the type locality in Carrizo Arroyo. 

Description. — Body small. Thoracomeres subequal, pleura apparently rounded, 
except eighth which appears posteriorly serrate. Pleomeres subequal, posterior comers 
of at least first 3 pleura serrate. Telson rectangular, developed distally with 2 sets of 
tooth-like spines, medial distal set larger than lateral proximal pair. Uropodal rami, 
blade-like and subequal, slightly longer than telson, exopod with distal tooth-like spines 
on lateral margin just anterior of where diaeresis might be, endopod margins finely 

Remarks. — Ox\\y one specimen allows any inference to be made concerning the 
anatomy of the animal. Two other specimens (SDSNH 29140) appear to preserve only 
part of the trunk segment series. The observed thoracopods are of such a diaphanous 
preservation as to preclude any more concrete conclusions about them other than that 
they seem to be equally developed back to and including the eighth pair. None of the 


Plate 1 

Figures A & B. Erythrogaulos carhzoensis new species, holotype, SDSNH 25 1 4 1 ; A) whole specimen, x 7; 
B) closeup of posterior abdomen and tailfan, note spines on posterior of pleura (p), spines on distal telson 
(t) and lateral margin of uropodal exopod (e), and setose margins of uropodal rami (arrows), x 19.6. 

Figures C-E. Acanthotelson stimpsoni Meek and Worthen, 1865; C & D) latex peels of holotype, X 346, 
X 1.7; E) syntype oi A. event Meek and Worthen, 1868, ISGS 3066 (made a junior synonym of A. stimpsoni, 
by Packard 1886), x 1.2. 


anterior limbs seemed specialized in any way, though these had to be partly destroyed 
in preparation in order to fully reveal the cephalon. No traces of pleopods were noted, 
and this, combined with the fact that thoracopods are clearly detectable and with the 
general small size of the body, might suggest the possibility, as with Minicaris brandi, 
that pleopods were either greatly reduced or not present on this species. 

The distinctively serrate pleura and spinose telson warrant separate generic status 
for this species from its nearest relative, Minicaris brandi of the Lower Carboniferous. 

The associated biota in the Red Tanks Member includes: numerous plants dom- 
inated by the gymnosperm genera Walchia and Cordaites, but also including Callipteris, 
sphenopsids, and lycopsids; a most diverse array of uniramians including insects and 
myriapods; the eurypterid Adelophthalmus luceroensis; ostracodes; brachiopods; and 
spirorbid worms. Kues and Kietzke (1981) interpret the paleoecology of the Carrizo 
Arroyo fauna as representing a fresh to brackish water habitat on a delta plain. The 
extreme delicacy of the preservation from this locality also indicates quick burial under 
anoxic conditions with little postdepositional disturbance. 

Family ACANTHOTELSONIDAE Meek and Worthen, 1865 

Diagnosis.— Thoracic exopods unisegmental and flap-like. Anterior thoracopods 
raptorial. Pleopods biramous and flap-like. 

Type genus.— Acanthotelson Meek & Worthen, 1865. 

Remarks. — brooks (1962a, b) chose to place the genera Acanthotelson and Uro- 
nectes in separate families based on the degree of raptorial development expressed in 
the anterior thoracopods. Although this is an important character, it is best utilized 
for distinction at the generic level. The unisegmental, flap-like nature of the thoracic 
exopods and rami of the pleopods in comparison to other palaeocaridacean families 
herein recognized warrants uniting all species with raptorial thoracopods into a single 

Genus ACANTHOTELSON Meek and Worthen, 1865 

Diagnosis. — First thoracopod markedly reduced. Second and third thoracopods 
raptorial. Telson and uropods styliform. 

Type species.— Acanthotelson stimpsoni Meek and Worthen, 1865. 

Acanthotelson stimpsoni Meek and Worthen, 1865 
Fig. 9; Plate 1, figs. C-E, Plate 2, fig. A 

v.* 1865 Acanthotelson stimpsoni Meek and Worthen, p. 47. 

V. 1866 Acanthotelson stimpsoni Meek and Worthen. Meek and Worthen, p. 401, pi. 32, figs. 6, 6a-f. 

1868a Acanthotelson eveni Meek and Worthen. Meek and Worthen, p. 27. 

V. 18686 Acanthotelson stimpsoni Meek and Worthen. Meek and Worthen, p. 549, 2 figs. 

V. 1868Z? Acanthotelson eveni Meek and Worthen. Meek and Worthen, p. 551. 4 figs. 

1880 Acanthotelson stimpsoni Meek and Worthen. Brocchi, p. 10, pi. 1, fig. 11. 

1884 Acanthotelson stimpsoni Meek and Worthen. White, p. 176, pi. 37, fig. 4-5. 

1 184 Acanthotelson eveni Meek and Worthen. White, p. 177, pi. 38, figs. 4-7. 

V. 1886<2 Acanthotelson stimpsoni Meek and Worthen. Packard, p. 123, pi. 1, figs. 1-3, pi. 2, figs. 1-3. 

1886a Acanthotelson eveni Meek and Worthen. Packard, p. 125. 

1890 Eileticus anthracinus Scudder, p. 420, pi. 38, fig. 5. 

1890 Eilecticus aequalis Scudder, p. 421, pi. 38, figs. 6-9. 

1896 Acanthotelson stimpsoni Meek and Worthen. Caiman, p. 799, pi. 2, fig. 16. 

1901 Acanthotelson species Fritsch, p. 74, fig. 398. 

1909 Acanthotelson stimpsoni Meek and Worthen. Smith, p. 575, fig. 62. 

1911a Acanthotelson stimpsoni Meek and Worthen. Caiman, p. 159. 

191 la Acanthotelson eveni Meek and Worthen, Caiman, p. 159. 

1916 Acanthotelson stimpsoni Meek and Worthen. Cockerell, p. 234. 

1916 Acanthotelson stimpsoni Meek and Worthen. Vanhoffen, p. 146, fig. 12. 

1916 Acanthotelson eveni Meek and Worthen. Vanhoffen, p. 148. 

1916 Acanthotelson species. Vanhoffen, p. 148, fig. 14. 

1919 Acanthotelson species. Pruvost, p. 85. 

7927 Acanthotelson stimpsoni Meek and Worthen. Chappuis, p. 605. 


Figure 9. A) Reconstruction of Acanthotelson stimpsoni, scale 5 mm (modified from Brooks 19626); B) 
lailfan to same scale as body; dorsal views of C) right antennule and D) antenna; E) posterior thoracopod. 
Appendages slightly enlarged. 

1931 Acanthotelson event Meek and Worthen. Van Straelen, p. 1 1. 

1931 Acanthotelson stimpsoni Meek and Worthen. Van Straelen, p. 12. 

1959 Acanthotelson event Meek and Worthen. Slewing, p. 2. 

1959 Acanthotelson stimpsoni Meek and Worthen. Siewing, p. 3. 

1962a Acanthotelson stimpsoni Meek and Worthen. Brooks, p. 236. 

V. 19626 Acanthotelson stimpsoni Meek and Worthen. Brooks, p. 230, pis. 55-59; Text-pl. 10, 11a. 

7965 Acanthotelson species Noodt, p. 83. 

1969 Acanthotelson stimpsoni Meek and Worthen. Brooks, p. R355, figs. 165-2, 171. 

1969a Acanthotelson stimpsoni 
1969b Acanthotelson stimpsoni 
1976a Acanthotelson stimpsoni 
1976b Acanthotelson stimpsoni 
1979a Acanthotelson stimpsoni 
1979b Acanthotelson stimpsoni 
1981a Acanthotelson stimpsoni 
1981b Acanthotelson stimpsoni 

Meek and Worthen. Schram, p. 219, Table 1. 

Meek and Worthen. Schram, p. 201. 

Meek and Worthen. Schram, p. 21. 

Meek and Worthen. Schram, p. 411. 

Meek & Worthen. Schram, p. 28, Table 1. 

Meek and Worthen. Schram, p. 167, fig. 1, Table 2. 

Meek and Worthen. Schram, p. 131, text-fig. 5b, Table 2. 

Meek and Worthen. Schram, p. 9, fig. in text. 

7952 Acanthotelson event Meek and Worthen. Kent, p. 15. 

Diagnosis. — Second joinX of antennal peduncle shorter than first or third. Telson 
styHform, equal to or slightly longer than uropods. 

Lectotype. — X 346 (Plate 1 , figs. C and D). Mazon Creek area, Will County, Illinois. 
Francis Creek Shale, Carbondale Formation (Westphalian C), Pennsylvanian. (Im- 
properly designated a holotype by Brooks \962b.) 

Other localities. — {Sqq Schram 1976a) Illinois State Geol. Surv. core T-4 (816 feet) 
NW V4, SW '/4, SE »/4, sec 25, T2S, R14W, Wabash County Ilhnois; Dykersburg Shale, 
Carbondale Formation, Pennsylvanian. Sec. 4, T9S, RIE, '/z mile west of Carterville, 
Williamson County; gray shale above #6 (Herrin) Coal, Brereton Cyclothem, Penn- 
sylvanian. Abandoned Chieftan Mine, 7 miles south of Terre Haute, Indiana, east of 
Highway 41; Lower Shelbum Formation, Pennsylvanian. 

Description.— CQXi^3\on with short rostrum, cervical and precervical grooves. Eyes 
small and stalked. Antennular peduncle 3-segmented, proximal and distal joints large, 
medial segment short, flagella well developed with inner branch shorter than outer 
branch. Antennal protopod with short proximal segment bearing nephropore and long 
distal segment, no scaphocerite, very long flagellum with proximal 2 joints peduncular. 
Antennules and antennae with setose inner peduncular margins. Mandible massive, 
palp well developed. Maxillule with 3-segment palp. Maxilla with at least proximal 
segment of palp large. 

First thoracomere reduced in length. Second through fourth thoracomeres pro- 
gressively longer than first. Last 4 thoracomeres subequal, last 3 thoracomeres have 


anterior margins with raised ridge. Thoracic pleura simple. First thoracopod reduced, 
possibly as short maxillipede. Second and third thoracopods biramous, endopods large, 
spinose, and raptorial in form. Five posterior thoracopods of ambulatory form, with 
epipodites, exopods of single segment (flap-like), endopods with short ischium and 
dactylus, and long merus, carpus, and propodus joints. 

Pleomeres similar in size to posterior thoracomeres, first through fifth pleura with 
posterio-ventral comers serrate, fifth and sixth pleomere posterior margins serrate. 
Sixth pleomere not elongate. Pleopods as biramous setose paddles. Telson as long spike, 
margins with alternating spines and setae. Uropodal rami as blades, margins with 
alternating spines and setae. Spikes or uropods and telson reinforced with median 

Remarks.— Though recognized as a distinct taxon since the time of Meek and 
Worthen (1865); a complete and reliable description and accurate reconstruction of .4. 
stimpsoni was not available until Brooks {\962b). However, actual photo illustrations 
of the type series of A. stimpsoni have not been prepared until now, except for the 
single exception of Brooks {\962b, pi. 54, fig. 4). Latex peels illustrated here of the 
lectotype, X346 (Plate 1, figs. C and D), are taken from the specimen which was used 
as the basis for one of the drawings in Meek and Worthen (1868/^:549, fig. B). A 
paralectotype, X3442, (Plate 2, fig. A) was the basis for another drawing in Meek and 
Worthen (1868^7:549, fig. A). The tail on X3442 was also apparently used by Meek 
and Worthen as an addition to augment their drawing (p. 551, fig. A) of ISGS 3066 
(Plate 1, fig. E). ISGS 3066 is also a syntype of another species, A. eveni, since syn- 
onymized (Packard 1886) with A. stimpsoni. 

Heretofore, understanding of what constitutes the genus Acanthotelson has been 
clouded incredibly by the incorrect use of the name Eileticus Scudder, 1882, by Eu- 
ropean workers. Eileticus {sensu stricto) is now generally conceded to be a myriapod. 
However, as pointed out by Brooks (1962Z):258), Scudder designated as a separate 
taxon, E. aequalis, what turned out to be a poorly preserved specimen of A. stimpsoni. 
Some European workers built upon this confusion and have applied the name Eileticus 
to other taxa that have turned out not to be Acanthotelson. For example, E. cf. aequalis 
of Pruvost ( 1 9 1 9) is probably Pleurocaris, and E. pruvosti Vandenberghe ( 1 960) is likely 
better placed in Nectotelson. Each of these cases is discussed in detail elsewhere, under 
the appropriate taxon designation. 

Acanthotelson kentuckiensis new species 


Frederick R. Schram and Donald Chesnut* 

*Kentucky Geological Survey, Lexington 

Fig. 10; Plate 2, figs. B-F 

Diagnosis. — ^QgmQnXs of antennular peduncles subequal. Telson subtriangular but 
long and narrow, shorter than uropods. 

//o/o/y/7£'.-SDSNH 23722 (Plate 2, fig. B). Black Oak Coal, Inc. strip mine, near 
Silverville, McCreary County, Kentucky, 2 miles north of Tennessee state line; 84°26'30" 
N, 36°38'42" W. Black fissile shale above River Gem Rider Coal, Lower Breathitt 
Formation, Middle Pennsylvanian. 

Other material. -SDSNH 23723 (Plate 2, fig. E), 23724, 23725. 

Descriptions. — Aniennules well developed, with 3 subequal segments (Plate 2, fig. 
F) composing the peduncle, about equal in size to peduncle of antennae. Antennal 
peduncular segments apparently short, no scaphocerite noted. 

All pleomeres about equal in length, last 3 with paired longitudinal dorsal ridges, 
dorsal posterior margin of sixth pleomere concave. Telson (Plate 2, figs. C & D) tri- 
angular in outline, narrow, with dorsal median ridge, margins furrowed and setose, 
distal setae more strongly developed. Uropodal rami styliform, each with reinforcing 
rib flanked by slight furrows, exopodal margins with strong setae (especially laterally), 
endopodal margins finely setose. 


,—■ -^ 

"l 1 ^' ~T~T~Ttt 


' / / ; ' ' ' ' > 


Figure 10. A) Diagrammatic rendition of what is currently known about the form of Acanthotelson ken- 
tuckiensis. scale 5 mm; B) tailfan to same scale as body. 

Remarks.— T\iQ most complete specimen (SDSNH 23722) lacks a thorax and all 
other specimens are of tailfans only. However, though anatomical information about 
this species is minimal, the distinctive nature of the tailfan, especially the styliform 
uropods, is so different from that seen in A. stimpsoni as to require, pending some 
future evidence to the contrary concerning thoracopods, a separate species for this 
material within the genus Acanthotelson. 

The shorter and broader telson of ^. kentuckiensis is more primitive than the long 
styliform tail oi A. stimpsoni. Apparently, the styliform expression is allometric since 
the smallest specimen of ^. kentuckiensis, SDSNH 23723, has the broadest and shortest 
telson (Table 1) while the larger specimens are narrower and longer. A. stimpsoni, by 
contrast, is a generally larger and more robust species than A. kentuckiensis, and has 
a very long, styliform telson. 

The biota associated with A. kentuckiensis indicates a fresh to brackish water facies. 
In addition to occasional fish scales and teeth, abundant remains were collected of the 
pelecypod Anthraconaia, and fossils of the plant Calamites were common. These ob- 
servations on the biota are reinforced by the nature of the black, fissile, canneloid shale 
in which the animals are found. The fossils occur at the base of a generally coarsening 
sequence of shales and sandstones, beginning with the carbonaceous shales with abun- 
dant fossils, and grading into an increasing arenaceous sequence with interbedded gray 
and black shales. These beds overlay another coarsening sequence with the River Gem 

Table 1. Measurements in mm on material of Acanthotelson kentuckiensis. Comparative data on telson 
measurements included for two representative specimens of A. stimpsoni. see text for discussion. * Ho- 


Length Max. 

Length A, Length Length width Ration 

cephalon peduncle a^ telson telson tl:tw 

Length Length 

uropodal uropodal 

exopod endopod 

A. kentuckiensis 

♦SDSNH 23722 

SDSNH 23723 

SDSNH 23724 

SDSNH 23725 

A. stimpsoni 

SDSNH 17454 
SDSNH 5210 






























. ■ttj '.-,•".. ."j< >•■ 


® ^^ 

_ J- 

Plate 2 

Figure A. Acanlhotelson stimpsoni Meek and Woilhen, 1865, paralectotype X 344-Z, x 1.3. 

Figures B-F. Acanthotelson kentuckiensisnev/ species; B-D, F) holotype SDSNH 23722; B) whole specimen 
displaying cephalon and abdomen, thorax missing, x3.9; C, D) closeup of tailfan counterparts, x7.8; F) 
closeup of cephalon, note antennular peduncle with 3 subequal segments (arrow). E) SDSNH 23723, note 
relatively wider telson (t) in relation to length than that seen in C or D, x6. 

Rider Coal at the base, on top of which are a siltstone and shale grading into a heavy 
burrowed argillaceous sandstone. 

Genus URONECTES Bronn, 1850 (=GAMPSONYCHUS Burmeister, 1855) 

Diagnosis. — No rostrum. First thoracomere moderately reduced. Second thora- 
copod raptorial. Telson and uropods broad and rounded, uropods with straight diaere- 
sis, broad tailfan formed from overlapping elements. 


Figure 11. A) Reconstruction of Uronectes fimbriatus, scale 5 mm (modified and corrected from Brooks, 
1962Z?); B) tailfan to same scale as body; C) dorsal view of right antenna slightly enlarged. 

Type species. — Gampsonyx fimbriatus Jordan, 1 847. 

Uronectes fimbriatus (Jordan), 1847 
Fig. 1 1 

*1847 Gampsonyx fimbriatus Jordan, p. 89, pi. 2. 
1848 Gampsonyx fimbriatus Jordan. Bronn, p. 575. 
7550 Gampsonyx fimbriatus Jordan. Bronn, p. 575. 
1850 Uronectes fimbriatus (Jordan). Bronn, p. 575. 

1854 Gampsonyx fimbriatus Jordan. Jordan & von Meyer, p. 1, pi. 2. 

1855 Gampsonychus fimbriatus (JoTdan). Burmeister, p. 191, pi. 10, figs. 12-14. 

1856 Uronectes fimbriatus (Jordan). Roemer, p. 202. 

1856 Gamsonychus fimbriatus (Jordan). Roemer, p. 202, p. 672. 

1873 Gampsonyx fimbriatus Jordan. Feistmantel, p. 593, pi. 18, figs. 9-11. 

1877 Carcinurus fimbriatus (Jordan). Goldenburg, p, 35, pi. 2, figs, lb, 2-7. 

1880 Gampsonyx fimbriatus Jordan. Brocchi, p. 10, pi. 10, fig. 7. 

1885 Gampsonychus fimbriatus (Jordan). Zittel, p. 672, fig. 857 (in part). 

1886b Gampsonychus fimbriatus (Jordan). Packard, p. 130, fig. 1. 

1896 Gampsonyx fimbriatus Jordan. Caiman, p. 798, pi. 2, fig. 17. 

1900 Gampsonyx fimbriatus Jordan. Eastman in Zittel, p. 659, fig. 1382. 

1901 Gampsonychus fimbriatus (Jordan). Fritsch, p. 72, pi. 159, text-fig. 377. 

1902 Uronectes fimbriatus (Jordan). Caiman, p. 66. 

1909 Gampsonyx fimbriatus Jordan. Smith, p. 568, fig. 53-55. 
1916 Gampsonychus fimbriatus (Jordan). Vanhoffen, p. 143, fig. 7-8. 
1927 Uronectes fimbriatus (Jordan). Haack, p. 733, 3 figs. 
1927 Uronectes fimbriatus (Jordan). Chappuis, p. 605. 
1931 Uronectes fimbriatus (Jordan). Van Straelen, p. 18. 

1958 Uronectes fimbriatus (Jordan). Malzahn, p. 355. 

1959 Gampsonychus fimbriatus (Jordan). Slewing, p. 1. 
1962(2 Uronectes fimbriatus (Jordan). Brooks, p. 236. 

1962^ Uronectes fimbriatus (Jordan). Brooks, p. 230, text-pl. 1 lb. 

1963 Uronectes species. Noodt, p. 82. 

1969 Uronectes fimbriatus Jordan. Brooks, p. R355, figs. 165-3, 173. 

1969a Uronectes fimbriatus Jordan. Schram, p. 221, table 1. 

7972 Uronectes species Jordan. Boy, p. 47, fig. 2. 

1974 Uronectes fimbriatus Jordan. Schram & Schram, p. 101. 

7979 Uronectes fimbriatus Jordan. Schram & Schram, p. 170. 

7952 Uronectes species Schneider, et al., p. 75, fig. 5. 

Diagnosis. — Sixih thoracomere somewhat enlarged over adjacent segments. Sixth 
pleomere long. 

Lectotype.—]k 4a, b, from the Kramer Ironworks of Lebach, near Saarbriicken, 
Saarland, West Germany. Rotliegende, Lower Permian. 


Paralectotype.—Jk 5. 

Other localities. — Pfeffelbach, near Kusel, Rheinlandpfalz, West Germany; Rot- 
liegende. Oberhof (Schweitzerhatte), near Zella-Mehlis, Thiiringia, East Germany; Ob- 
erhofer Beds, Lower Permian. 

Diagnosis. —Cephalon with faint cervical groove, no rostrum. Antennule peduncles 
3-segmented, proximal segment very long, distal 2 joints short, flagella moderately 
developed. Antennal protopod with 2 subequal segments, scaphocerite oval, flagellum 
moderately long with proximal 2 segments peduncular. 

First thoracomere moderately reduced. Thoracic pleura simple, with slight furrow 
along margins. Sixth thoracomere somewhat longer dorsally than others. Second thor- 
acopod large, spinose, and raptorial. Second through eighth thoracopods ambulatory, 
more or less subequal, ischium very short, merus through propodus moderate, dactylus 
very small. 

First 5 pleomeres with finely serrate posterior margins, pleura acuminate anteriorly 
with slight furrow on margins. Sixth pleomere elongate. Telson rounded, margins setose. 
Uropods flap-like, margins setose, exopod with straight diaeresis and reinforced with 
lateral thickened rib. 

Remarks.— The reconstructions of U. fimbriatus prepared by Brooks (1962^?, 1969) 
generally reflect an accurate view of the creature, except for the fact that he mistakenly 
drew 7 abdominal segments instead of 6 (corrected here in Fig. 1 1). The description 
in his text indicates the proper number. 

The classic Lebach locality has been the source of U. fimbriatus specimens in 
museums around the world. The freshly collected material was a black shale. The 
characteristic red rock with white fossils developed only after the specimens were 
"roasted" at the Kramer Ironworks, driving off" the volatile organics in the shale and 
fossils, and leaving a calcitic residue behind on a rock residue high in siderite. Specimens 
from other localities were found by me while searching various European collections. 
The Staatssamlungen flir Palaontologie in Munich has an "unroasted" specimen from 
the Rothegende (BS 1975 I 164) from Pfefffelbach, near Kusel, not too far from Lebach. 
The Munich collection also has a specimen from the Oberhofer Beds from near Oberhof, 
in Thiiringia (BS 1953 XXVIII 21) in a strange 3-dimensional preservation. The Pa- 
laontologisches Museum of the Museum fiir Naturkunde in Berlin also has specimens 
(PMB A. 62-67) identified as U. fimbriatus from Thiiringia (H.-E. Gruner, pers. comm.), 
as does the San Diego Natural History Museum (these a gift of Dr. J. Schneider of the 
Bergakademie, Freiberg). 

Uronectes kinniensis Schram and Schram, 1979 

Fig. 12 

v.* 1979 Uronectes kinniensis Schram and Schram, p. 169, pi. 1, text-fig. 1. 
1981a Uronectes kinniensis Scharm and Schram. Schram, p. 133, text-fig. 4g. 

Diagnosis. — YourXh. thoracomere moderately reduced; eighth thoracomere with 
lateral semicircular ridges. Fifth pleomere elongate. 

Holotype.-\]^NM 235625. Kinney Clay Pit, SE 'A, Sec. 18, T9N, R6E, Bernalillo 
County, New Mexico. Madera Formation, Virgilian, Pennsylvanian. 

Description.— CephaXon apparently undecorated, no rostrum. Antennal protopod 
of 2 subequal segments, scaphocerite subtriangular and setose, at least proximal-most 
joint of flagellum peduncular. 

First and fourth thoracomeres moderately reduced. All thoracomeres except eighth 
undecorated, pleura simple. Eighth thoracomere with small, lateral, paired, semicircular 
ridges. Thoracomeres 3 through 8 subequal (details obscure). 

Pleopods with finely serrate posterior margins. Fifth pleomere elongate. Telson 
rectangular, rounded distally, and apparently distally serrate. Uropods as broad flaps, 
exopod with straight diaeresis, at least endopods setose. 

Remarks.— The lack of knowledge about the first and second thoracopods makes 
it difficult to place this species in Uronectes without any hesitation. However, as orig- 


Figure 12. A) Reconstruction of Uronectes kinniensis, scale 5 mm (modified from Schram and Schram 
1979); B) tailfan to same scale as body; C) dorsal view of right antenna slightly enlarged. 

inally reported (Schram and Schram 1979:170) the overall aspects of the anatomy 
(especially the serrate pleomere margins, lack of rostrum, moderately reduced first 
thoracomere, and straight diaeresis) come closest to Uronectes, and U. kinniensis is 
best left within that genus for the time being. 

Genus PALAEOSYNCARIS Brooks, \962b 

Diagnosis. — First thoracomere very reduced, second thoracomere moderately re- 
duced. Second and third thoracopods raptorial. Telson oval, with spinose margins. 
Uropodal exopods laterally spinose. 

Type species.— Palaeosyncaris dakotensis Brooks, \962b. 

Palaeosyncaris dakotensis Brooks, \962b 
Fig. 13; Plate 3, figs. B-E 

v.* 1962^ Palaeosyncaris dakotensis Brooks, p. 251; pi. 65, figs. 3, 4, pi. 66; text-pl. 14, fig. a. 
1969 Palaeosyncaris dakotensis Brooks. Brooks, p. R355, figs. 169-2, 170-2. 

Figure 13. A) Reconstruction of Palaeosyncaris dakotensis, scale 5 mm (corrected from Brooks 1962^); 
B) tailfan to same scale as body; C) dorsal view of right antenna slightly enlarged. 


Plate 3 

Figure A. Palaeosyncaris micra new species, holotype, PE 2496, x4.1. 

Figures B-E. Palaeosyncaris dakotensis Brooks, 1 962*; B-D) holotype, USNM 1 43409; B) whole specimen. 
x2.4; C) closeup of lailfan, x7; D) closeup of anterior thoracopods, note the inflated meri on thoracopods 
(2) and (3) and antennal scale (arrow), x 8; E) counterpart of holotype, Univ. of North Dakota collection, 
note inflated meri on thoracopods (2) and (3) and antennal scale (arrow), x5.5. 

1969a Palaeosyncaris dakotensis Brooks. Schram. p. 216. table 1. 
1974 Palaeosyncaris dakotensis Brooks. Schram and Schram, p. 95. 

Diagnosis.— A\\ segments with transversely striate decoration. Abdominal pleura 
with posterior margins serrate. 

Holotype. — \]SNM 14309 (Plate 3, figs. B-D) (counterpart, unnumbered, in col- 
lection of University of North Dakota, Plate 3, fig. E). Borehole Casimer Duletski No. 


1,81 70-8 1 80 feet NW 'A, NW 'A, Sec. 1 6, T 1 39N, R99W, Stark County, North Dakota. 
Heath Shale, Upper Mississippian. 

Description. — Eyes small, eye stalk long. Antennules with 3-segment peduncle, 
proximal joint equal to distal 2 joints, flagella well developed. Antennae with small, 
oval, finely setose scaphocerite, proximal 2 joints of flagellum peduncular witfi median 
margins setose, flagellum well developed (but of undetermined length). 

First thoracomere greatly reduced, second thoracomere moderately reduced; sec- 
ond and all other somites (Plate 3, fig. B) with transverse striae, pleura rounded, and 
ventral margins with furrow. First thoracopod apparently reduced; second through 
eighth thoracopods robust, merus somewhat longer than other subequal joints, merus 
on second and third thoracopods (Plate 3, figs. D-E) inflated (possibly spinescent), 
dactylus long and pointed, second and third thoracopods raptorial. Eighth thoracic 
pleuron posteriorly extended and margin serrate. All tergites with marginal furrows, 
especially prominent on pleura. 

Abdominal pleura anteriorly somewhat rounded, posteriorly pointed with mar- 
gins serrate. Last pleomere somewhat elongate. Telson long, oval, and marginally spi- 
nose, terminal median spines reduced in comparison to adjacent members of series. 
Uropodal exopod laterally spinose, endopod margins finely setose (Plate 3, fig. C). 

Remarks. — Though obviously well-preserved thoracic epipodites and exopods ap- 
pear not to have been present on the type specimen, there is some indication on the 
coxa of the third thoracopod of USNM 143409 of a foramen for an epipodite. This 
same appendage may also preserve part of an exopod arising from the basis. 

The thoracopodal endopods of this species are all strongly developed. The inflated 
meri on the second and third thoracopods may well have been capable of acting like 
subchelae in opposition to the carpi on these appendages, which appear to be proximally 
narrow and with a rather disto-posterior spiniform crest. In this regard USNM 143409 
appears to have partially preserved the sockets of articulating spines on the merus of 
the second thoracopod disto-posteriorly. 

Brooks (1962) compared P. dakotensis to Praeanaspides praecursor, mainly on the 
basis of similarities of tergal ornament. The tailfans, however, are now known to be 
quite different (Schram 1 979a). Furthermore, the identification of raptorial thoracopods 
on P. dakotensis would appear to ally this species with members of the Acanthotel- 
sonidae. Placement in this family should not be without query, however, since complete 
knowledge of the thoracic exopods and pleopods would be necessary before unques- 
tioned affiliation could be sanctioned. Pleopods are not preserved on the type coun- 

The other syncarid of the Heath Shale is Squillites spinosus. This latter species is 
collected from "paper shale" outcrops of the Heath in central Montana, while the rock 
of the North Dakota core which contains P. dakotensis is a well-indurated, blocky, 
black shale. The associated fauna on the core section with P. dakotensis is largely 
composed of partially pyritized cyzicoid branchiopods and casts of indeterminate os- 

Palaeosyncaris micra. new species 
Fig. 14; Plate 3, fig. A; Plate 4, figs. A-E 

Diagnosis. — 'Qody small. Segments mooth, undecorated. Abdominal pleura not 
serrate. Telson with spinose setae increasing in size distally. 

Holotype. — PE 2496 (Plate 3, fig. A). Mazon Creek area; Will, Crundy, and Kan- 
kakee Counties, Illinois. Francis Creek Shale, Carbondale Formation (Westphalian C- 
D), Pennsylvanian. 

Description. — Body small, tergites smooth. Cephalon with short rostrum. Anten- 
nules with 3 subequal joints in peduncle. Antenna with small setose scaphocerite. 

First thoracomere greatly reduced, second moderately reduced, third through eighth 
subequal with anterior comers rounded (Plate 4, figs. A, C). First thoracopod reduced, 
about one-half the length of ambulatory thoracopods. Second and third thoracopods 


Figure 14. A) Reconstruction of Palaeosyncaris micra, scale 5 mm; B) tailfan to same scale as body; C) 
dorsal view of right antenna slightly enlarged. 

robust, raptorial, dactyli with well-developed terminal spines (Plate 4, fig. A). Third 
through eighth thoracopods ambulatory; meri and carpi long, ischia, propodi, and 
dactyli short; exopods flap-like (Plate 4, fig. C). 

Abdominal pleura anteriorly and posteriorly reduced. Sixth pleomere somewhat 
longer than anterior pleomeres. Telson oval (Plate 4, fig. D), medial margins with 
spinose setae, with setae increasing in size distally. Uropods (Plate 4, figs. D, E) setose, 
setae of lateral margin of exopod spinose, exopod reinforced with medial rib and 
apparently possessing an oval diaeresis. 

Remarks. — ThQ small size and generally incomplete preservation of these fossils 
has resulted in their being mistakenly sorted by previous workers (including myself) 
as "small and poorly preserved" examples of the other two Mazon Creek syncarids, 
Acanthotelson stimpsoni and Palaeocaris typus. We are indebted to Mr. Stephen L. 
May for recognizing these specimens as a separate species, and bringing it to our 
attention. Representative measurements are given in Table 2. 

The establishment of a third Mazon Creek syncarid now brings the crustacean 
assemblage of the brackish water biotope in the American Pennsylvanian into accord 
with that of the European Carboniferous (Schram 1981a). Both faunas now have a 
pygocephalomorph associated with 3 species of syncarid (see Table 3). However, there 
does not appear to be a point-for-point analogy between the syncarid species. The 
American faunas have 2 rapacious acanthotelsonids and 1 palaeocarid, whereas the 
British fauna syncarids are in apparently 3 different families. Palaeosyncaris micra 
occurs in both the Essex and Braidwood faunas of Johnson and Richardson (1966). 

Table 2. Representative measurements in mm of species of Palaeosyncaris. * Holotypes. 









P. dakotensis 

*USNM 143409 









P. micra 

*PE 2496 







PE 11670 






PE 1268 





PE 12174 








PE 37912 







PE 37915 














- «» 

'-e- - 

<*-^ - 


■u-,. ^ 





Plate 4 

Figures A-E. Palaeosyncaris micra new species; A) latex peel of holotype, PE 2496. closeup of anterior 
end, note the raptorial thoracopods 2 and 3 (arrows), and the reduced first thoracomere (1), x9.8; B) latex 
peel of PE 11670, with rounded thoracic and abdominal pleura, x4.5; Q latex peel of PE 37915, note 
progressively increasing lengths of anterior thoracomeres (1-3) and large flap-like exopods (arrows), x 7; D) 
latex peel of PE 12168, showing spination on telson, x 10.7; E) latex peel of PE 12174, showing spines on 
lateral margin of uropodal exopod and possible diaeresis (arrow), setation of endopod, and all but distal- 
most portions of the telson, x 7.4. 

Table 3. Crustaceans found in Late Carboniferous brackish water habitats in North America and Europe. 

Mazon Creek Faunas 

Westphahan British Coal Measures 


Acanthotelson stimpsoni 

Pygocephalus cooperi 


Acanthotelson stimpsoni 
Palaeocaris typus 
Palaeosyncaris micra 

Praeanaspides praecursor 
Palaeocaris retractata 
Pleurocaris annulatus 

Family PALAEOCARIDIDAE Meek and Worthen, 1865 

Diagnosis.— Thovdic'xc exopods flap-like, pleopods annulate. 

Type genus.— Palaeocaris Meek and Worthen, 1865. 

Remarks. — The genus Paleocaris has long been treated as a catchall taxon for every 
incompletely known Paleozoic eumalacostracan which has not had any evident cara- 
pace. This has been complicated by the fact that Palaeocaris itself was incorrectly 
understood, in the sense of Brooks (1962). Unfortunately, much remains to be discov- 
ered about this most important genus of Paleozoic syncarids; however, the diagnostic 
combination of annulate pleopods with flap-like thoracic exopods provides a focus 
upon which further work in the group can be based. It is possible the genera Brook- 
syncaris, Palaeorchestia, and Williamocalmania may belong to this family, but these 
fossils yet lack relevant information about the appendages to allow definitive placement 
of them into the Palaeocarididae. 

Genus PALAEOCARIS Meek and Worthen, 1865 

Diagnosis. — M.3.n6\h\QS massive, first thoracomere greatly reduced. Sixth pleomere 
posterior margin deeply concave. Uropodal rami margins very setose, exopod with 
pronouced diaeresis, exopod distinctly longer than endopod, endopod distinctively 
longer than telson. Telson oval, margins bearing stout setae. 

Palaeocaris typus Meek and Worthen, 1865 

[=Acanthotelson inaequalis Meek and Worthen, 1865] 

Fig. 15; Plate 5, figs. A-C 

1865 Acanthotelson inaequalis Meek and Worthen, p. 48. 
v.* 1865 Palaeocaris typus Meek and Worthen, p. 49. 

1866 Acanthotelson inaequalis Meek and Worthen. Meek and Worthen, p. 403, pi. 32, fig. 7. 
V. 1866 Palaeocaris typus Meek and Worthen. Meek and Worthen, p. 405, pi. 32, figs. 5, 5a-d. 
1868(3 Palaeocaris typus Meek and Worthen. Meek and Worthen, p. 28. 

18686 Palaeocaris typus Meek and Worthen. Meek and Worthen, p. 552, figs. 1, 2. 

1880 Palaeocaris typus Meek and Worthen. Brocchi, p. 9, pi. 1, figs. 8-10. 

1884 Palaeocaris typus Meek and Worthen. White, p. 179, pi. 38, figs. 1-3. 

V. 18866 Palaeocaris typus Meek and Worthen. Packard, p. 129, pi. 7, figs. 1-2. 

1889 Palaeocaris typus Meek and Worthen. Packard, p. 213. 

1896 Palaeocaris typus Meek and Worthen. Caiman, p. 796, pi. 2, fig. 15. 

7909 Palaeocaris typus Meek and Worthen. Smith, p. 570. text-figs. 56-58. 

79/6 Palaeocaris typus Meek and Worthen. Vanhoffen, p. 141, fig. 5. 

7976 Acanthotelson inaequalis Meek and Worthen. Vanhoffen, p. 147. 

7927 Palaeocaris typus Meek and Worthen. Chappuis, p. 605. 

V? 1957a Palaeocaris species. Copeland, p. 595; pi. 6, fig. 5. 

V? 1957b Palaeocaris of. typus Meek and Worthen. Copeland, p. 47; pi. 15, fig. 1. 

7959 Palaeocaris typus Meek and Worthen. Siewing, p. 3. 

1959 Acanthotelson inequalis Meek and Worthen. Siewing, p. 102. 

7967 Palaeocaris typus Meek and Worthen. Rolfe, p. 548. 

V. 19626 Palaeocaris typus Meek and Worthen. Brooks, p. 240, pis. 60-64, text-pis. 12 (fig. a), 13. 

7965 Palaeocaris species Noodt, p. 82. 

1969 Palaeocaris typus Meek and Worthen. Brooks, p. R348, figs. 165-1, 167, 170- la, 171. 

1969a Palaeocaris typus Meek and Worthen. Schram, p. 219, table 1. 

1969b Palaeocaris typus Meek and Worthen. Schram, p. 201. 

7972 Palaeocaris typus Meek and Worthen. Secretan, p. 3. 



Figure 15. A) Reconstruction oi Palaeocaris typus, (corrected from Brooks 1962^?), scale 5 mm; B) tailfan 
to same scale as body; dorsal views of right C) antennule and D) antenna; E) posterior thoracopod. Appendages 
slightly enlarged. 

797^ Palaeocaris typus Meek and Worthen. Schram and Schram, p. 101. 

1976a Palaeocaris typus Meek and Worthen. Schram, p. 21. 

1976b Palaeocaris typus Meek and Worthen. Schram, p. 411. 

7975 Palaeocaris typus Meek and Worthen. Schminke, p. 235, fig. 17. 

1979a Palaeocaris typus Meek and Worthen. Schram, p. 28, table 1. 

1979b Palaeocaris typus Meek and Worthen. Schram, p. 167, table 2. 

1981a Palaeocaris typus Meek and Worthen. Schram, p. 131, text fig. 5c, table 2. 

1981^ Palaeocaris typus Meek and Worthen. Schram, p. 9, fig. in text. 

Diagnosis. — HQdid to thorax ratio 1:4. Scaphocerite longer than the two peduncular 
segments of antennal flagellum. Sixth thoracomere larger dorsally than any other tho- 
racomere. Uropodal diaeresis slightly curved to straight, outer margin of exopod with 
widely spaced spinose setae along its length terminating in 3 spines just anterior to 
diaeresis. Telson ovoid but wider proximally than distally. 

Holotype. — X33S (Plate 5, fig. A), Mazon Creek area, Will County, Illinois. Francis 
Creek Shale, Carbondale Formation (Westphalian C), Pennsylvanian. [Brooks (\962b: 
248) states that the types of this species "are misplaced or lost." However, close 
comparison of X338 with the description of Meek and Worthen (1868, fig. A:552), 
and the more detailed treatment in their original description (Meek and Worthen 1 865), 
reveal that this specimen was undoubtedly the basis for these texts and the 1868 
illustration, and is thus almost certainly the holotype.] 

Other localities.— Abandoned Chieftan Mine, 7 miles south of Terra Haute, In- 
diana, east of Highway US-4 1 ; Lower Shelbum Formation, Pennsylvanian. Abandoned 
strip mine talus 1.8 miles west of Windsor, Missouri, on Highway MO-2. 

Description.— Cephalic shield smooth, except for slight lateral groove at level of 
mandible. Rostrum small. Optic notch prominent. Ventral margins of cephalic shield 
rounded and whole. Eyes moderate in size, stalk with prominent (peracarid-like) papilla. 
Antennular peduncle 3-segmented, proximal-most joint as long as distal 2 joints, medial 
margins setose, inner flagellum shorter than outer flagellum, outer flagellum about one- 
third body length. Antennal protopod with short proximal joint and large distal joint, 
scaphocerite oval and setose, 2 basal flagellar joints peduncular, medial margins of 
peduncular joints setose, flagellum equal to body length. Mandible massive, promi- 
nently projecting below cephalic shield margin, palp of at least 2 segments. Maxillules 
and maxillae with palps (details uncertain). 

First thoracomere markedly reduced, sixth thoracomere larger than any other. 
Thoracic pleura broadly rounded anteriorly, posterior margin straight. First thoracopod 


Plate 5 

Figures A-C. Palaeocaris typus Meek and Worthen, 1 865; A) latex peel of holotype, X 388, x 1 .9; B) YPM 
19765, showing the flap-like thoracopodal exopods, x 4.5; C) PE 23237. showing annulate pleopods (arrows), 

Figure D. Palaeocaris retracta Caiman, 1932, I 13971, with short section of an annulate pleopod (arrow), 

about one-half the size of succeeding appendages. Thoracopods 2 through 8 subequal, 
epipods present (details not known), exopods broadly flap-like (Plate 5, fig. B) and 
setose, endopods with short ischia and dactyli, meri and carpi long and subequal. 
propodi one-half the length of carpi. 

Pleomeres with posterior margins finely setose, pleura of first 5 abdominal segments 
as in thoracomeres. Pleopods annulate (Plate 5. fig. C) [not flap-like as reported by 
Brooks 1962]. Telson oval, somewhat wider anteriorly than medially or posteriorly, 
margins with strong setae. Uropods flap-like, faint median reinforcing rib on setose 


Figure 16. A) Reconstruction of Palaeocaris retractata, (modified from Schram 1979a), scale 5 mm; B) 
tailfan to same scale as body; C) dorsal view of right antenna; D) thoracopod. Appendages slightly enlarged. 

rami, exopods with lateral margin spinose, setae distally developed as 3 small spines 
just anterior of straight to slightly curved diaeresis. 

Remarks. — brooks (1962Z?) made some errors in anatomical interpretation of the 
P. typus specimens available to him. He felt the antennal peduncle had 5 segments, 2 
protopodal and 3 flagellar. The mistake arose in interpreting a preservation anomaly 
on the distal protopodal joint, mistaking a longitudinal crack in that joint for a lon- 
gitudinal suture. 

More importantly. Brooks reconstructed the pleopods of P. typus as flap-like, and 
compared them to those of Acanthotelson stimpsoni. The pleopods are rarely well- 
preserved on P. typus. The thoracic exopods are clearly flap-like (e.g., YPM 19765, 
Plate 5, fig. B). The pleopods, however, are annulate [YPM 19731, YPM 19765, PE 
23237 (Plate 5, fig. C), PE 37893, PE 37957, PE 37976]. It is the correction of our 
understanding of this feature and its detection on other species of Palaeocaris that 
delineates the family Palaeocarididae from other palaeocaridacean syncarids. 

Palaeocaris retractata Caiman 1932 
Fig. 16; Plate 5, fig. D; Plate 6, fig. A 

V. 191 Id Palaeocaris praecursor (Woodward). Caiman, p. 488, figs. 1, 2a, 3. 

V. 1914 Palaeocaris species. Peach, p. 146, pi. 4, fig. 9. 

*1932 Palaeocaris retractata Caiman, p. 541. 

1959 Palaeocaris retractata Caiman. Siewing, p. 101. 

V. 1961 Palaeocaris retractata Caiman. Rolfe, p. 546, pi. 68, fig. 8, text-fig. 1. 

1962b Palaeocaris retractata Caiman. Brooks, p. 248. 

1979(2 Palaeocaris retractata Caiman. Schram, p. 106, figs. 50, 51. 

1979b Palaeocaris retractata Caiman. Schram, p. 170, table 2. 

1981a Palaeocaris retractata Caiman. Schram, p. 131, text fig. 5e, table 2. 

1982 Palaeocaris retractata Caiman. Schram, p. 123, fig. 8. 

Diagnosis. — Head to thorax ratio 1:2.8. All thoracic segments subequal. Uropodal 
diaeresis a rounded to sigmoid curve, outer margin of exopod armed with spines. Telson 
ovoid, margin with spinose setae. 

Holotype. — In 29012. Clay Craft open works, Cosely near Dudley, Worcestershire. 
Ten foot Ironstone Measures, Lower Similis-Pulchra Zone, Middle Coal Measures. 

Other locality. — WqsI flank Bilberry Hill, in Lickey Hills southwest of Birmingham, 
Warwickshire; Keele Beds, Weslphalian D. 

Plate 6 

Figure A. Palaeocaris retractata Caiman, 1932, I 13973, showing setose margins of telson, x7. 

Figures B-G. Palaeocaris secretanae new species: B) holotype, AM 7423, x 3.6; C) AM 7243, closeup of 
cephalon, showing stalked eyes, bases of antennules and antennae, and mouthparts, x 10.7; D) AM 4293, 
closeup of annulate pleopod, x 22.3; E) AM 7861, showing optic notch, cephalic groove, large mandible, and 
thoracopodal exopods, x 10.7; F^ AM 5019, showing cephalic groove and the markedly reduced first tho- 
racomere, xio.7; G) AM 7810, lateral preservation of posterior abdomen, with telson (t) and pleomeres 
(numbered), x 10.7. 

Figure H. Nectotelson /cre/"c/7 (Fritsch) 1875, B 77621, tailfan, x6.6. 



Descriptions. — Cephalic shield smooth, slight lateral groove at level of mandible, 
premandibular portion of cephalon as long or longer than the posterior region. Rostrum 
small. Optic notch slight. Eyes small to medium in size. Antennular peduncle with 3 
subequal joints, the most proximal with an optic fossa. Antennal protopod with distal 
joint long, scaphocerite small and oval, 2 basal-most joints of flagellum petiuncular. 
Mandible large with prominent incisor process. 

All thoracomeres, except the markedly reduced first, subequal in length. Second 
through fourth thoracic pleura subquadrangular, posterior pleura broadly rounded an- 
teriorly, all pleura with slightly marginal furrows. Second through eighth thoracopods 
with epipods possibly flap-like, moderate flap-like exopods, endopodal joints subequal 
with a tendency to shorten as one proceeds distally. 

Second through sixth pleomeres with setose posterior margins. Pleopods annulate 
(Plate 5, fig. D). Telson oval, margins setose (Plate 6, fig. A). Uropodal exopod with 
rounded to sigmoid diaeresis, lateral margins spinose, other margins of rami setose (at 
least distally). 

Remarks.— Reexamination and preparation of available material. In 29013, In 
29014, and especially I 13971 (Plate 5, fig. D), indicates that the pleopods of/', retractata 
are annulate. This was not noticed in the redescription of Schram {\919a). 

Now that other species of Palaeocaris are better understood, the large cephalon 
(small head to thorax ratio) is seen as quite diagnostic for this species. Other species, 
P. typus and P. secretanae, have relatively smaller heads. 

Restudy in July of 1980 of all P. retractata material mentioned in Schram (1979<3) 
aflowed me an opportunity to reconsider the identity of doubtful specimens in light of 
these collateral studies of all fossil syncarids. I now feel that an incomplete specimen, 
GSL RAE 1 29 1 , is not an example of P. retractata. This correction does not affect our 
understanding of the anatomy of this species. However, it does shorten the biostratig- 
raphic range of P. retractata (Schram 1979(3:7, fig. 1), now understood to extend only 
from Westphalian B to D, i.e., from the Lower Similis-Pulchra Zone up into the Tenuis 

Palaeocaris secretanae new species 
Fig. 17; Plate 6, figs. B-F; Plate 7, figs. A & B 

V. 1980a palaeocarid syncarid. Secretan, p. 24, pi. 1. 

V. 1980a ?ceratiocarid phyllocarid. Secretan, p. 28, pi. 2, figs. 1, 2. 

V. 1980a ?eocaridacean. Secretan, p. 28, pi. 3, figs. 2, 3. 

V. 1980a pygocephalomorph. Secretan, p. 30, pi. 3, figs. 5, 6. 

V. 1980a ?palaeostomatopod. Secretan, p. 30. 

V. 1980a "specimens enigmatiques." Secretan, p. 32, pi. 4, figs. 2 & 6. 

1980b Palaeocaris. species. Secretan, p. 414, pis. 1-4, fig. 1. 

1981 Palaeocaris cf. P. retractata Caiman. Pacaud et al., p. 40. 

1982 Palaeocaris cf. P. retractata Caiman. Rolfe et al., p. 426. 

Diagnosis. — Head to thorax ratio 1:4.6. All thoracic segments subequal. Uropodal 
diaeresis markedly circular, outer margin of exopod armed with small spines distally 
near diaeresis, margins of rami with long dense setae, rami reinforced with heavy 
median ribs. Telson ovoid with distal end blunt, margin with long spines. 

Holotype.— AM 7423-24 (Plate 6, figs. B & C). From shales above First Blanzy- 
Montceau Coal (=-Puits St. Louis), Stephanian B, Upper Carboniferous. St. Louis open 
cast mine, Montceau-les-Mines, France. 

Etymology. — Named in honor of Dr. Sylvie Secretan, who first recognized the 
nature and significance of this material, and who has been a major figure in organizing 
and coordinating the scientific study of the important biota of the Montceau-les-Mines 

Description. — Cephalic shield smooth, slight lateral groove at level of mandible 
(AM 5019; 7861, Plate 6, figs. E & F). Rostrum small to moderate in size. Optic notch 
prominent (AM 7861). Eyes moderate to large (AM 7423-24). Antennular peduncle 
with 3 subequal segments, median margins setose (AM 6 1 37-38), flagella well developed 


Figure 17. A) Reconstruction of Palaeocaris secretanae, scale 5 mm; B) tailfan to same scale as body; 
dorsal views of C) antennule and D) antenna; E) thoracopod. Appendages slightly enlarged. 

(but of undetermined length). Antennal protopod with median margins setose (AM 
7794), scaphocerite oval, small in size and setose, 2 proximal-most joints of flagellum 
peduncular and very large (AM 7524). Mandible (AM 5744) with large incisor process 
and well-developed palp. 

Thoracic and abdominal pleura with gently rounded anterior comers and well- 
developed posterior comers (AM 7810, Plate 6, fig. G), all body segments (except first) 
subequal. First thoracopod reduced in size (AM 5019). Second through eighth thora- 
copods with long cylindrical epipodites (AM 7861), exopods large and flap-like with 
setose margins, endopodal ischium and dactylus small, merus and propodus long, and 
carpus moderate in length (AM 7423-24). 

Annulate pleopods setiferous (AM 4293, Plate 6, fig. D; 4377, 7424). Telson with 
margin spinose (AM 7436, Plate 7, fig. B). Uropodal rami spatulate (AM 5080, Plate 
7, fig. A), reinforced with strong median ribs, exopodal lateral margin with small spines 
distally (AM 5080), diaeresis strongly circular (Plate 7, fig. B), ramal margins with 
dense array of long setae. 

Remarks.— Though closely resembling P. retractata, P. secretanae is easily distin- 
guished by its short cephalon (shorter than that of any species of Palaeocaris) and the 
spatulate nature of the uropodal rami, i.e., more narrow proximally than distally. 

Although the general preservation of most of the material of P. secretanae is 
exceptionally fine, those specimens that were not so well-preserved can be rather con- 
fusing to interpret. This accounts for the variety of tentative assignments made by 
Secretan (1980a, b). Her non-syncarid identifications are all based on poorly preserved 
specimens. In point of fact, though the biota at Montceau-les-Mines is among the most 
diverse in the Carboniferous, P. secretanae remains the only malacostracan presently 
known from that fauna, save for one specimen that is possibly a phreatoicid isopod. 
Measurements comparable to those made by Brooks (1962b) on P. typus are provided 
in Table 4. 

Family SQUILLITIDAE Schram and Schram, 1974 

Diagnosis. — Thoracic exopods annulate, pleopods annulate and either uni- or bi- 

Type genus. — Squillites Scott, 1938. 



V'*^ '♦> 


Plate 7 
Figures A & B. Palaeocaris secretanae new species; A) AM 5080, tailfan, x 10.7; B) AM 7436, tailfan, 

Figures C-G. Nectotelson krejcii (Fritsch) 1875; C) B 7762b, showing prominent cephalic groove (arrow) 
and 8 subequal thoracomeres (numbered), x 5.6; D) M 1 042, typical form and preservation of Czech localities, 
X 7; E) B 7762i, showing subequal trunk segments (numbered) and fragments of annulated pleopods (arrows), 
X 7; F) B 7762d, latex peel showing antennae and thoracopodal endopods, x 6.7; G) M 1033, showing setose 
margins of telson, x5. 


Table 4. Representative length measurements in mm of species of Palaeocaris* Holotype. 









P. secretaneae 

AM 3424 



































































































































P. typus X 








P. retractata x 





Remarks.— TogQXhtx the annulate thoracic exopods and pleopods of these species 
most resemble those seen in the living Anaspididae within the order Anaspidacea. 
However, the first thoracomere is not fused to the cephalon in these fossils: equal in 
size to all other thoracomeres in Nectotelson, slightly reduced in Squillites, and greatly 
reduced in Praenaspides. 

Genus SQUILLITES Scott, 1938 

Diagnosis. — ¥'\rs,\ thoracomere only slightly reduced. Uropods narrow, spatulate, 
and setose. Telson subtriangular, armed with moveable spines. 
Type species.— Squillites spinosus Scott, 1938. 

Squill it es spinosus Scott, 1938 
Fig. 18 

v.* 1938 Squillites spinosus Scott, p. 508, figs. 1, 2. 

1939 Squillites spinosus Scott. Berry, p. 467. 

1962^ Squilites spinosus Scott. Brooks, p. 254, pi. 53, figs. 1, 3; text-pl. 14, fig. d. 

1965 Squillites species. Noodt, p. 82. 

7967 Squillites spinosus Scott. Secretan, p. 173, fig. 8. 

1969 Squillites spinosus Scotx. Brooks, p. R355, figs. 169-1 and 170-3. 

1969a Squillites spinosus Scott. Schram, p. 216, table 1. 

V. 1974 Squillites spinosus Scott. Schram and Schram, p. 96, pis. 1-2, text-figs. 1, 2. 


Figure 18. A) Reconstruction of Squillites spinosus, (modified from Schram and Schram 1974), scale 5 
mm; B) tailfan to same scale as body; C) dorsal view of right antenna slightly enlarged. 

J 97 9a Squillites spinosus Scott. Schram, p. 114. 

1981 Squillites spinosus Scott. Schram, p. 133, text-fig. 4. 

7952 Squillites spinosus Scott. Schram, p. 122. 

Diagnosis. — Since there is but one species, the diagnosis of the species is the same 
as that of the genus. 

Holotype. — Xl 2 19. One-half mile south of Heath, Fergus County, Montana; Heath 
Shale, Big Snowy Group, Upper Mississippian. 

Description. — Stalked compound eyes small and spherical. Antennule with 3-seg- 
ment peduncle, flagella long. Antennal protopod with only 1 segment observed, scaph- 
ocerite oval and setose, very long flagellum with proximal 2 segments large and pe- 
duncular. Cephalon with marked broad rostrum, lacking any cervical grooves, pair of 
semicircular mid-dorsal ridges. 

Thoracomeres with 4 anterior pleura medially pointed and 4 posterior pleura 
rounded, pair of semicircular ridges mid-dorsal on each thoracomere, first thoracomere 
slightly shorter than others. Thoracopods subequal, ischia short and equal to bases, 
meri long, carpi through dactyli short. 

Pleomeres variously decorated, first through fifth with setose posterior margins, 
first and second with mid-dorsal paired semicircular ridges, and third through fifth with 
large immobile posteriorly directed spines. First pleuron rounded, second through fifth 
pleura with posterior comers denticulate. Setose pleopods robust and uniramous. Sixth 
pleomere elongate. Uropods as oval flaps, margins finely setose. Telson subtriangular, 
with median keel and 1 7 pairs of moveable marginal spines. 

Remarks.— There is a slight reduction in the size of the first thoracomere, but 
reconsideration of the original material of Schram and Schram (1974) leaves some 
question as to whether their first thoracopod is as well developed as those on the other 
thoracic segments. 

The apparent single joint on the antennal protopods may be an artifact of pres- 
ervation, but oddly coincides with an apparent similar phenomenon on Praeanaspides 
praecursor, which, if it is confirmed, may provide another derived feature to characterize 
the family. 

Genus PRAEANASPIDES Woodward, 1908 

Diagnosis. — ¥\rs\ thoracomere very reduced. Pleopods biramous. Uropodal exo- 
pod with distinct circular diaeresis. Telson rectangular, laterally spinose. 


Figure 19. A) Reconstruction of Praeanaspides praecursor, scale 5 mm (redrawn from Schram 1979(2); B) 
tailfan to same scale as body; C) dorsal view of right antenna slightly enlarged. 

Type species.— Praeanaspides praecursor Woodward, 1 908. 

Praeanaspides praecursor Woodward, 1 908 
Fig. 19 

v.* 1908 Praeanaspides praecursor Woodv/ard, p. 385, figs. 1-5. 

V. 1908 Paleocaris lansboroughi Peach, p. 55, pi. 8, figs. 8-10. 

1911 Paleocaris lansboroughi Peach. Woodward, p. 363. 

7927 Paleocaris praecursor {V^ood\^ard). Chappuis, p. 605. 

1959 Praeanaspides praecursor Woodward. Siewing, p. 10. 

1959 Paleocaris praecursor V^oodvjavd. Siewing, p. 101. 

1932 Paleocaris praecursor (V^oodward). Caiman, p. 537, figs. 1, 2. 

1962b Paleocaris praecursor (Woodward). Brooks, p. 249. 

1969a Paleocaris praecursor (Woodward). Schram, p. 220, table 1. 

7976 Praeanaspides praecursor Woodward. Schram, p. 411. 

1979a Praeanaspides praecursor Woodward. Schram, p. 112, figs. 54, 55. 

1979b Praeanaspides praecursor Woodward. Schram, table 2. 

7957 Praeanaspides praecursor Woodward. Schram, p. 131, table 2. 

Diagnosis. — S>mcQ there is but one species, the diagnosis of the species is the same 
as that of the genus. 

Lectotype.—GSh 30213-14. Shipley Hall, I'A miles NW of Ilkestone, Derbyshire, 
England. Clay Ironstone, top Modiolaris Zone, Middle Coal Measures. 

Other locality.— GvQQnhxW, or Woodhill Quarry, near Kilmaurs, Ayrshire; roof of 
Pinnies Main Coal, Middle Coal Measures, Westphalian B. 

Description.— A.r\\Qr\n\x\Qs with 3-segment peduncle, middle joint shorter than either 
proximal or distal unit, flagella relatively short. Antennae with only single segment 
visible in protopod, oval non-setose scaphocerite, flagellum very long with proximal 2 
joints peduncular. Cephalon unomamented, slight rostral projection. 

First thoracomere smooth with no ornament, all other segments with 3 or 4 tergal 
ridges and rounded pleura. First thoracopod apparently reduced (never seen). Second 
through eighth subequal, ischium and merus longer than carpus and propodus (dactyls 
not seen). Pleopods with exopod slightly longer than endopods, rami thin. Uropodal 
exopod lateral margin spinose, endopod subtrapezoidal with its longer margin medial. 
Telson rectangular but somewhat bilobed terminally, lateral margin with 12-13 pair 
of moveable spines. 


Figure 20. A) Reconstruction of Nectotelson krejcii, scale 5 mm; B) tailfan to same scale as body; C) dorsal 
view of right antenna; D) posterior thoracopod. Appendages slightly enlarged. 

Remarks. — ThQ apparently single segment antennal protopod is possibly an artifact 
of preservation, a short proximal segment just may not be observable on known material 
(however, see remarks on 5". spinosus). The same problem applies to the thoracopodal 
dactyli; they were probably short and simple, but were simply not preserved on any 
specimens now available. 

Of all the palaeocaridaceans, P. praecursor comes closest to resembling the anas- 
pidid anaspidaceans in regard to its very reduced first thoracic segment and in the 
character of its thoracic and abdominal appendages. 

Genus NECTOTELSON Brocchi, 1880 

Diagnosis. — A\\ thoracopods (?) and thoracomeres subequal. Pleopods biramous. 
Uropods spatulate, diaeresis circular. Telson oval and spinose. 
Type species. — Gampsonychus krejcii Fritsch, 1875. 

Nectotelson krejcii (Fritsch) 1875 
Fig. 20; Plate 6, fig. H; Plate 7, figs. C-G; Plate 8, fig. A-D 

1870 Gampsonychus species, Fritsch, p. 34. 

1873 Gampsonychus fimbriatus Jordan. Feistmantel, p. 593, pi. 18, figs. 9-12. 

v.* 1875 Gampsonychus krejcii Fritsch, p. 104, fig. 265. 

V. 1880 Nectotelson rochei Brocchi, p. 10, pi. 1. 

1885 Gampsonychus fimhriatus (Jordan). Zittel, p. 672 (in part). 

1885 Nectotelson rochei Brocchi. Zittel, p. 673. 

V. 1901 Gasocaris krejcii (Fritsch). Fritsch, p. 66, figs. 371-376, pis. 156-158. 

790/ Nectotelson rochei Brocchi. Fritsch, p. 74. 

1909 Gasocaris krejcii (Fritsch). Smith, p. 572, figs. 59-61. 

1919 Gasocaris species Fritsch. Pruvost, p. 85. 

1919 Nectotelson species Brocchi. Pruvost, p. 85. 

1931 Gasocaris species Fritsch. Van Straelen, p. 5. 

1959 Gasocaris krejcii (Fritsch). Siewing, p. 5. 

1959 Nectotelson rochei Brocchi. Siewing, p. 103. 

? 1960 Eileticus pruvosti Vandenberghe, p. 690, fig. 2, pi. 17. 

1965 Nectotelson species Noodt, p. 82. 

1965 Gasocaris species Noodt, p. 82. 


• '''^- ^-JR*^- 15^V ^^ ** ' > ■•-V:■'■■■ 

Plate 8 

Figures A-D. Nectotelson krejcii (Fritsch) 1875: A) lectotype, M 1050, x4.5; B) Me 40, showing diaeresis 
on uropodal exopod (arrow), x5; C) B 7762k, latex peel showing cephalon (c) and thoracopodal epipodites 
(arrows), x7; D) B7762J, showing antennules and antennae, and thoracopods, x7. 

Figure E. Pleurocaris annulatus Caiman, 1911; I 13814, latex peel showing thoracic pleura, tergite deco- 
ration, and proximal portions of thoracopods, x6.2. 

1969 Palaeocaris krejcii {¥hX's,c\\). Brooks, p. R355, figs. 170- lb. 
1969 Palaeocaris rochei (QtoccYlx). Brooks, p. R355, figs. 170-lc. 
1969a Palaeocaris krejcii (Fritsch). Schram, p. 221, table 1. 
1969a Palaeocaris rochei (Brocchi). Schram, p. 221, table 1. 
1972 Nectotelson rochei Brocchi. Secretan, p. 1, 1 fig. 


Diagnosis.— Since but a single species is known, the diagnosis of the species is the 
same as that of the genus. 

Lectotype. — M 1050 (Plate 8, fig. A). Humboldt Mine, Nyfan, near Pilsen, Bo- 
hemia. Gaskohle, Lower Permian. 

Other localities. — FriXsch (1901) also recorded this species from the Gaskohle of 
the Krimitz Mine and in Tremosna. Brocchi (1880) described the same species under 
another name from the Lower Permian shales near Autun, central France. 

Description.— Cephalon lacks prominent rostral extensions (M 1035, B 7762g, 1), 
marked by rather deep and prominent mid-dorsal (B 7762b, Plate 7, fig. C) groove 
parallel to posterior margin which shallows as it extends toward ventral margin (B 
7762h). Eye oval, moderate in size (B 7762j, Plate 8, fig. D). Antennular peduncles 
subequal, 3-segmented middle joint slightly shorter than other two (M 1035, B 7762b, 
e & j, Plate 8, fig. D. Antennal protopod with short proximal segment and longer distal 
joint (B 7762e, B 7763b), these with slight longitudinal ridges (Me 40). Scaphocerite 
short, oval, and setose (Me 40, B 7762e), overlapping 2 proximal-most peduncular 
joints of flagellum. Antennular and antennal flagella well developed, [of undetermined 
length because of lack of preservation of distal joints (M 1042, Plate 7, fig. D; B 11626. 
& j, Plate 7, fig. F)]. Median margin of antennal peduncle marked by row of short 
denticulae (B 7762e, B 7763k). 

Mandible large but not apparently heavily sclerotized or mineralized (B 7763a), 
with an incisor process (B 7762i, B 7763c). Maxillules and maxillae with small palps, 
maxillary palp seems to have several segments (B 7763a). 

First thoracomere large (B 7762i, Plate 7, fig. E), with fine serrations on posterior 
margin (B 7763k). Thoracic pleura rounded posteriorly (M 1050, B 77621, and the TV. 
rochei holotype). First thoracopod large [but not completely preserved on either the 
Prague or Paris specimens (e.g., B 7763g)], with moderately long proximal unit (?is- 
chium) followed by short merus, long carpus, and distal to the knee may be at least 
another short unit. Second through eighth thoracopods (Plate 7, fig. F) subequal; with 
small epipods (B 7762k, Plate 8, fig. C); stout annulate exopods (M 1042, B 7762d & 
k); short coxae, bases, and ischia; long meri; and distal to knee progressively shorter 
carpi, propodi, and dactyli (M 1042 in part, B 7762e, B 7763f & g); posterior margins 
of endopods with apparently fine setae (B 7762a). 

Pleomeres with acute postero-ventral comers (M 77631); second through sixth 
pleomeres with finely serrate posterior margins (M 42, M 1033, M 1054, B 7762b). 
Sixth pleomere almost twice as long as any other segment (M 1054, B 7762f, B 7763b). 
Pleopods with robust, annulate rami subequal (M 1054, B 7762b & m, B 7763), and 
setose (M 42). 

Uropods longer than telson; protopod short, with 2 faint longitudinal ridges, spat- 
ulate rami (B 77621, Plate 6, fig. H), densely setose, and reinforced with sclerotized 
ribs along most of their lengths (M 1044, M 1054, B 7763h, A^. rochei type). Exopods 
laterally spinose (M 1054, B 7762a, B 7763e) diaeresis circular (Me 40, Plate 8, fig. B), 
segment beyond diaeresis as a narrow oval (M 1033, B 7763d). Telson elongate and 
oval; ornamented with stout, short, moveable spines set in sockets (M 1033, Plate 7, 
fig. G; M 1044, M 1054, B 7762b & m, B 7763e). 

Remarks. — Both the Bohemian and French specimens are preserved as carbonized 
films, with varying degrees of pyrite replacement, on a fine-grained thinly-bedded black 
shale. The pyrite, especially on the Bohemian material, is highly reflective, and makes 
studying and photographing specimens extremely difficult. In only a few instances were 
fossils preserved as external molds, and these tend to occur on shale fragments from 
Autun with a higher content of clay mineral. The matrix in these latter cases was 
cemented enough to allow one latex peel to be made, but otherwise the shale and fossils 
are too friable to allow such treatment. In general, the French material is somewhat 
better preserved in contrast to the Czech specimens. 

There are subtle differences between the two suites of specimens, especially as 
regards the antennal peduncles. That described above generally represents that seen on 
the better preserved French specimens. The Bohemian material seems to have a some- 
what shorter distal segment of the protopod, a somewhat longer scale, and no detectable 


Table 5. Representative measurements in mm of Nectotelson krejcii. 








B 7762b 





B 7762c 





B 7762g 


B 7762h 


B 77621 



B 77621 


B 7763a 



B 7763b 


B 7763c 









B 77631 


B 7763J 


B 7763k 


B 77631 


M 1033 


M 1035 








medial denticulae. The Czech specimens also have slightly more distinctive spines 
along the pleomere margins. However, in light of all the other fine and detailed points 
of agreement between the two series of specimens, and caveats due to vagaries of 
preservation, I feel that A^. rochei Brocchi must be synonymized with A^. krejcii (Fritsch). 
Nor, in light of the distinctive form of the thoracic exopods and the first thoracomere, 
can this species be maintained in the genus Palaeocaris in the sense of Brooks (1969). 
Rather, A^. krejcii is more closely aligned with the Squillitidae. 

I found no evidence of any of the sexual dimorphism reported by Fritsch (1901: 
70-71). Representative measurements of specimens are given in Table 5. 

A vexing problem in this study has been what to do with Eilecticus pruvosti 
Vandenberghe, 1960. The published paper contains no reference as to where the types 
were deposited; the photographs, while good enough to be intriguing, are not of sufficient 
quality to facilitate detailed study; and the description of the specimens is too vague 
to be of any real help. Attempts to find the specimens or locate Dr. Vandenberghe have 
proved futile. However, certain items in the published description seem to hint that 
this material may belong within the genus Nectotelson. Vandenberghe related that E. 
pruvosti had an abdomen of 7 segments, a thorax of 7 segments, and the first thoracic 
segment fused with the cephalon. Counts of body segments on the published photo- 
graphs indicate 14 subequal segments between the head and tailfan. Thus segment 
count for the abdomen must be wrong, and it seems logical to infer that the abdomen 
had 6 segments and the thorax 8. The description also makes reference to a deep groove 
on the cephalon, and spatulate uropodal rami. All these features would seem to cor- 
respond to identical characters noted above in Nectotelson. Consequently, I choose to 
assign this species, with a query, to A^. krejcii. The question mark can be removed only 
by rediscovery and study of the type, or recollection at the original locality. 

E. pruvosti occurs at the top of the Griiner Group, near the Middle and Upper 
Stephanian boundary. The material came from a borehole in the Saint-Etienne basin, 
and is associated with the limuline Pringlia demaistrei, insects, fish scales, and the 
plant Odontopteris pseudoschotheimi. 

Family uncertain 

Remarks.— The familial taxonomy of the Palaeocaridacea being adopted here is 
an attempt to establish a more natural system than any used heretofore, and is based 
on comparative morphology of thoracic and abdominal appendages. Such a system, 


however, requires that certain features of the anatomy of these fossils be known before 
famiHal assignments can be made. Unfortunately, fossils do not always preserve all the 
features that one would like to have information about. The fossil syncarids are no 
exception to this, and as a result there are some Paleozoic taxa that cannot be placed 
within a family with any degree of certitude, though we can recognize them as distinct 
and valid genera and species. It was felt here that the issue should not be forced, and 
that it was preferable to simply recognize the uncertainty and treat these taxa as presently 
"unassignable." Thoughts as to their affinities can sometimes be offered, but it seems 
better to patiently await future data which will allow someone to definitively place 
these problematica. 

Genus PLEUROCARIS Caiman, 1911a 

Diagnosis.— Cephalon small, 2 cephalic grooves not joined laterally. Tergites dec- 
orated with lateral ridges. Thoracic pleura very large. Telson and uropodal rami styli- 

Type species.— Pleurocaris annulatus Caiman, 1911a. 

Pleurocaris annulatus Caiman, 1911a 
Fig. 21; Plate 8, fig. E 

? 1881 Palaeocaris burnetii Woodward, p. 534, pi. 14, figs. 3a, 3b. 

v.* 19 11a Pleurocaris annulatus Caiman, p. 156, fig. 1. 

V. 191 \b Pleurocaris annulatus Caiman. Caiman, p. 494, fig. 5. 

.1912 Eileticus cf. aequalis Scudder. Pruvost, p. 66, pi. 2, figs. 6, 7. 

79/5 Pleurocaris annulatus Caiman. Chappuis, p. 173. 

1919 Pleurocaris annulatus Caiman. Pruvost, p. 86, fig. 21, 22; pi. 25, fig. 11. 

.1919 Eileticus cf. aequalis Scudder. Pruvost, p. 89, fig. 23; pi. 25, fig. 12. 

1922 Eileticus aequalis Scudder. Pruvost, p. 149. 

1923 Pleurocaris annulatus. Caiman. Pruvost, p. 149. 
7927 Pleurocaris annularis Caiman. Chappuis, p. 605. 
7959 Palaeocaris burnetii Woodward. Slewing, p. 101. 
7959 Pleurocaris annulatus Caiman. Slewing, p. 103. 
1962a Pleurocaris annulatus Caiman. Brooks, p. 236. 
7965 Pleurocaris species. Noodt, p. 83. 

1969 Pleurocaris annulatus Caiman. Brooks, p. 355, fig. 169-5, 172. 

1969a Pleurocaris annulatus Caiman. Schram, p. 220, table 1. 

? 1969a Palaeocaris burnetii Woodward. Schram, p. 220, table 1. 

7976 Pleurocaris annulatus Caiman. Schram, p. 411. 

1979a Pleurocaris annulatus Caiman. Schram, p. 103, figs. 48, 49. 

1979b Pleurocaris annulatus Caiman. Schram, p. 167, table 2. 

1981(2 Pleurocaris annulatus Caiman. Schram, p. 131, text-fig. 5f; table 2. 

Diagnosis. — ^mcQ there is but one species, the diagnosis of the species is the same 
as that of the genus. 

Holotype. — In 29008. Clay Croft mine, Coseley, near Dudley, Worcestershire, 
England. Ten foot Ironstone Measures, Lower Similis-Pulchra Zone, Middle Coal 
Measures, Upper Carboniferous. 

Other localities. — Pit no. 9, near Lens, Belgium; Black shale of the Insect beds, 
beneath the "veine Girard," Edouard Group, Westphalian C. Pit no. 4, Vicoigne Mines, 
France; Black shale, top of the "veine du Nord," Olympe Group, Westphalian A. 

Descriptions.— Cepha\on short, rostrum small. Stalked compound, eyes small and 
rounded. Details of antennules and antennae uncertain. 

First thoracomere short, about half the length of any other. All thoracomeres with 
2 laterally directed ridges on tergites; pleura very large, rounded, set off from tergites 
as lappets. Second through eighth thoracopods with well-developed endopods, meri 
long, carpi short (other joints indeterminable). 

Pleomeres decorated dorsally with lateral ridges as thoracomeres, pleura decrease 
in development posteriorly. Telson styliform, margins with 5 pair moveable spines. 
Pleopods possibly flap-like (I 14449). Uropods as blades; exopod straight, serrated 
laterally and less conspicuously so medially; endopod curved mediad, lateral margin 
faintly spined, medial margin distinctly so. 


Figure21. A) Reconstruction of P/eMrocamawwMtoiw, (modified from Caiman 1 9 1 1 a; and Schram 1979a), 
scale 5 mm; B) tailfan to same scale as body. 

Remarks. One specimen, In 14449, appears to have some poorly preserved prox- 
imal portions of the pleopods. These appear to be flap-like. However, they are of such 
quality as to be almost impossible to photograph. This was not noticed in the original 
study of Schram (1979a). Though the few British specimens known of this species 
(Schram 1979<3:121) are for the most part moderately well preserved, our knowledge 
of this taxon suffers because there are so few examples of it. All continental European 
material attributable to this species has been lost. However, examples of this species 
on the continent have apparently never been common, just as in Britain. 

Confusion in the identification of specimens of this species with Acanthotelson 
stimpsoni arises from the somewhat similar syliform telson and uropods. However, the 
short cephalon, subequal second through eighth thoracomeres, large thoracic pleura, 
and tergal decoration clearly justify a separate generic status for this species (Plate 8, 
fig. E). However, exact familial affinities must remain uncertain until such time as the 
structure of the thoracic exopods, distal joints of the endopods, and pleopods can be 
ascertained. The general form of the tailfan noted here, as well as a supposed analogue 
correspondence to A. stimpsoni in the Carboniferous brackish water habitat community 
(Schram 1981a) may suggest that P. annulatus could be eventually assigned to the 

Pruvost (1912, 1919) described 2 syncarid specimens which he variously referred 
to Eileticus cf. aequalis and/or Pleurocaris annulatus. The specimens were deposited 
in the museum at the University of Lille, but are now lost. The published descriptions 
and illustrations of these specimens, combined with Pruvost's own stated reservations 
on what he called E. cf. aequalis, indicate these were indeed examples off. annulatus. 


Diagnosis. — ¥'\rs\ thoracomere markedly reduced, second thoracomere longer than 
first but less than any other thoracomeres. Thorax shorter than abdomen, ratio about 
0.9:1. Telson elongate, subtriangular, distal end rounded. Uropodal rami oval, some- 
what longer than telson. 

Type species. — Palaeocaris vandergrachti Pruvost, 1922. 

jE'/.vwo/o^. — Named in honor of W. T. Caiman, among whose many accomplish- 
ments was his expertise on fossil and recent syncarids. 

Remarks.— The distinctive pattern of reduction of the anterior thoracomeres, the 
unique thorax-abdomen ratio, and the characteristic tailfan clearly separate this species 


Figure 22. A) Reconstruction of Williamocalmania vandergrachti, scale 5 mm; B) tailfan to same scale as 

from those herein included in the newly redefined genus Palaeocaris. Consequently, a 
separate generic designation is necessary for this taxon. Though obviously distinct from 
Palaeocaris or any other known fossil syncarid, the lack of sufficient information about 
body appendages dictates an uncertain family affinity for this species. 

Williamocalmania vandergrachti (Pniyost) 1922 
Fig. 22; Plate 9, figs. A-C 

v*1922 Palaeocaris vandergrachti Pruvost, p. 147, fig. 1. 

7927 Palaeocaris vandergrachti Pruvost. Chappuis, p. 605. 

1930 Palaeocaris vandergrachti Pruvost. Pruvost, p. 181, fig. 5, pi. 8. 

1959 Palaeocaris vandergrachti Pruvost. Slewing, p. 101. 

1969a Palaeocaris vandergrachti Pruvost. Schram, p. 220, table 1. 

Diagnosis. — Since but one species is known, the diagnosis of the species is the 
same as that of the genus. 

Lectotype.—NB 7183 Ech. no. 1 (see Pruvost 1930, plate 8, fig. la). Woensdrecht 
borehole (1 164-1 167 m). The Netherlands; Chokier Ampelite, Lower Namurian. 

Description. — Body moderate to large in size. Antennular peduncle large, with 3 
subequal segments. Antennal peduncles large, scaphocerite appears large and ovoid, 2 
(or 3) proximal-most segments of flagellum peduncular and very large. Cephalon with 
rostrum, prominent optic notch. 

First thoracomere reduced and closely associated with cephalon, second thorac- 
omere shorter than posterior thoracomeres but larger than first, all other thoracic and 
anterior abdominal segments subequal. Pleura somewhat subtriangular, attenuated along 
anterior margin (Plate 9, fig. C), with slight marginal furrows. Thoracopods apparently 
equally developed, exopods possibly flap-like (Plate 9, fig. A). Thorax somewhat shorter 
in length than abdomen. 

Sixth pleomere elongate. Telson long, subtriangular with distal end rounded, mar- 
gins with stout spinose setae (Plate 9, figs. B & C). Uropodal rami oval; exopod rein- 
forced proximally with medial rib, with slightly curved diaeresis, somewhat longer than 

Remarks.— The later treatment by Pruvost (1930) of this species is superior in 
most respects to the original description (Pruvost 1922), especially in regards to the 
plate figures which illustrate all 4 of the available specimens. Pruvost, however, claimed 
in his original description to have studied 1 2 specimens. The only substantially complete 




Plate 9 

Figures A-C. Williamocalmania vandergrachti (Pruvost) 1922; A) NB 7183 Ech. no. 3, note reduced first 
thoracomere (arrow), x7.6; B) NB 7183 Ech. no. 4, showing long, subtriangular telson, x7; Q NB 7183 
Ech. no. 2, showing somewhat acute abdominal pleura, x7.8. 

Figures D & E. Palaeorchestia parallela (Fritsch) 1876, part and counterpart of lectotype, CGH 593, 
showing the characteristic parallel-sided, distally circular telson; D) under water x3.6; E) x2.9. 


Table 6. 

Measurements of lengths in 


of Williamocalmania vandergrachti. 

* Lectotype. 






NB 7183 Ech 










* 2.3 








specimen now available, NP 7183 Ech. no. 1, clearly is the basis for the 1922 and 1930 
figure drawings, and thus designated here as the lectotype. Some measurements are 
provided in Table 6. 


Diagnosis. — First thoracomere only slightly reduced, sixth through eighth thorac- 
omeres slightly larger than second through fifth. Thoracomeres each with 2 transverse 

Type species.— Palaeocaris canadensis Brooks, 1962^7. 

Etymology.— '^2iVC\Qd. in honor of H. K. Brooks. 

Brooksyncaris canadensis (Brooks), \962b 
Fig. 23; Plate 10, fig. A 

v*1962^ Palaeocaris canadensis Brooks, p. 248; pi. 15, figs. 1, 2. 
1969a Palaeocaris canadensis Brooks. Schram, p. 220, table 1. 

Diagnosis. — Since there is but one species, the diagnosis of the species is the same 
as that of the genus. 

Holotype. — MCZ 5435 (Plate 10, fig. A); Confluence of Diligent and Ramshead 
Rivers, south of Diligent River, Cumberland County, Nova Scotia; Riversdale Group, 
Westphalian A. 

Description. -Cephsdon short (cephalon to thorax ratio 1:5.4); prominent cephalic 
groove; small postcephalic groove extending in arc dorsad from posterior margin. 

// // // 
'/ II ,1 


Figure 23. Diagrammatic rendition of what is currently known about the form oi Brooksyncaris canadensis, 
scale 5 mm. 


Antennual and antennal peduncles well developed [too poorly preserved to discern 

First thoracomere not much reduced. Second through eighth thoracopods ambu- 
latory, with large(?) epipodites. 

Remarks.— The specimens discussed by Copeland (1951 a, b) and referred by Brooks 
(1962/)) to this species have been reexamined by me. I concur with Copeland's original 
treatment of that material and refer those specimens to Palaeocaris cf. typus. Thus, the 
only material that is referable to B. canadensis is the holotype. 

Brooks ( 1 962^) described the thoracopods of this species as bearing epipodites. I 
have concurred with this for the time being, but it is difficult to clearly discern whether 
these structures are epipodites or flap-like exopods. These features are located very 
close to the base of the limbs, and I would also assume that the exopods probably 
would have been as poorly preserved as the endopods. However, the question remains 

The new genus is required because this species obviously does not belong in 
Palaeocaris as now understood, the latter taxon being characterized in part by the 
extreme reduction of the first thoracomere. However, none of the pertinent features of 
the thoracopods, abdomen, pleopods, or tailfan are preserved on the holotype that 
would allow us to place this species in any of the known genera of palaeocaridaceans, 
let alone family. Thus the establishment of a separate genus seems prudent. 

Genus PALAEORCHESTIA Zittel, 1885 

Diagnosis.— AnXennuXar peduncles smaller than those of antennae. Telson dis- 
tinctly rectangular. 

Type species. — Gampsonychus parallelus Fritsch, 1876. 

Remarks. — DQlerminrng the proper name for this genus poses a classic problem 
in untying the twisted strands of available names among Paleozoic syncarids. Jordan 
(1847) originally described Gampsonyx fimbriatus, unaware of the fact that the name 
of this genus was preoccupied in a bird, Gampsonyx swainsoni Vigors, 1825. Bronn 
(1850) did detect the synonymy and suggested the name Uronectes be applied to G. 
fimbriatus; while Burmeister (1855) independently caught the same synonymy and, 
unaware of Bronn's work, suggested the use of the name Gampsonychus for G. fim- 
briatus. Subsequent authors, until Chappuis (1927), ignored Bronn and used either 
Gampsonyx (and thus also ignoring the synonym) or Gampsonychus. 

In this context Fritsch (1876) described a new species Gampsonychus parallelus 
and allied it to G. fimbriatus. Zittel (1885) recognized the distinctive generic status of 
this species from fimbriatus and erected a new combination Palaeorchestia parallela. 
Brooks (1969) seemingly concurred with the Zittel distinction of/*, parallelus from 
what by then was known as U. fimbriatus, but implied that the proper generic assignment 
of the species was supposedly with Palaeocaris when he synonymized Palaeoorchestia 
{lapsus calumni), along with other syncarid genera, with Palaeocaris. Brooks was 
mistaken, since the taxa in question are distinctly different. One might be technically 
entitled to return to the use of Gamsonychus, since the use of Gampsonychus in Fritsch 
(1876) is not affected by any subjective synonymy in Burmeister (1855). However, to 
do so would be to: 1) return to the 19th and early 20th century confusion over the use 
of Gamsonyx-Gampsonychus, 2) minimize the importance of Zittel's initial recognition 
of the separate generic status of the type species, and 3) overlook Brooks' implicit 
acceptance of Zittel's work. For these reasons, I think the decisions of Brooks andZhtel 
should prevail and the name Palaeorchestia be used. 

Palaeorchestia parallela (Fritsch), 1876 
Fig. 24; Plate 9, figs. D & E 

1859 cf. Gampsonychus fimbriatus. Krejci, p. 79. 

v*1876 Gampsonychus parallelus Fritsch, p. 4, pi. 3 fig. 1, pi. 4. 

1885 Palaeorchestia parallela (Fritsch). Zittel p. 673, fig. 858. 


Figure 24. Dorsal reconstruction of what is currently known about the form of Palaeorchestia parallela, 
scale 5 mm. 

7907 Palaeorchestia parallela (Fritsch). Fritsch, p. 73. 
7976 Palaeorchestia parallela (Fritsch). Vanhoffen, p. 146, 
7959 Palaeorchestia parallela (Fritsch). Slewing, p. 103. 
7965 Palaeorchestia species Noodt, p. 82. 
7969 Palaeocaris parallela (Fritsch). Brooks, p. R355. 

fig. 11. 

Diagnosis. —SmcQ but one species is known, the diagnosis of the species is the 
same as that of the genus. 

Lectotype.—CGYi 593 (Plate 9, figs. D & E), from Lisek, northwest of Beraun, 
Bohemia, Czechoslovakia. Rodnitz Horizon, Coal Measures, Lower Permian. [The 
stratigraphic horizon is somewhat vague in the literature. One specimen in the British 
Museum (Natural History), In 35327, which resembles the preservation seen in the 
Czech types, is marked "Carboniferous, Nirzan, near Pilsen, Bohemia— Old Colin." 
seems to indicate the type Lower Permian area in Czechoslovakia.] 

Description. — ^ody moderate in size. Antennular peduncle with 3 subequal seg- 
ments, medial margin of second segment spinose, flagella well developed. Antennal 
protopod with short proximal segment, scaphocerite large and setose, basal joints of 
flagellum peduncular with distal segment twice as long as proximal. No rostrum. 

First thoracomere somewhat shorter than any other body segment. Thorax length 
more than one and one-half times that of abdomen. 

Sixth pleomere very long, with faint lateral groove about mid-length. Telson long, 
subrectangular, marginally setose. Uropodal rami spatulate, with strong median ribs, 
margins finely setose, exopod larger than endopod, endopod equal to or shorter than 
telson, exopod with circular diaeresis. 

Remarks. — ¥v\Xsc\\ (1876, 1901) referred to only one specimen. CGH 593 is ob- 
viously the basis of Plate 4 in his 1876 paper, and is thus designated the lectotype here. 
However, one additional specimen, CGH 592, is in the collections of the National 
Museum in Prague. It confirms the form of the head and antennae noted on the type, 
and clearly reveals the slight reduction in size of the first thoracomere. 

Aside from the antennules, antennae, and uropods, virtually nothing is known 
concerning any of the other appendages. CGH 593 preserves some remnants of anterior 
thoracopods. Both specimens are preserved in dorsal view in a brownish-gray mudstone, 
and details on the thorax and anterior abdomen are almost totally lacking. The addi- 
tional specimen in the British Museum, In 35327, contributes nothing towards un- 
derstanding this species. 



Figure 25. A) Partial reconstruction of Clarkecaris brasilicus, scale 5 mm; B) tailfan to same scale as body; 
dorsal views of C) right antennule and D) antenna slightly enlarged. 

Genus CLARKECARIS Mezzalira, 1952 

Diagnosis.— C^phdiXon with well-developed groove. Eight free thoracic segments 
short. Abdominal segments long, especially the sixth, pleura styliform. Telson subtrian- 
gular, with a narrow bifid terminus. 

Type species. — Gampsonyx brasilicus Clarke, 1920. 

Remarks.— Tht higher taxonomic placement of this species has been rather per- 
ipatetic. It was originally placed among the "gamsonychids," but when Mezzalira (1952) 
recognized the separate generic status of this creature he assigned it to the Uronectidae. 
However, Mezzalira's understanding of "uronectids" was not that of Brooks (1962a). 
The latter placed Clarkecaris in its own family within the anaspidaceans; though later 
(Brooks 1969) he reassigned it to the stygocaridaceans, still within its own family. 
Stygocaridines have since been recognized as a group within the Anaspidacea. However, 
reexamination of the types and an additional specimen studied and illustrated by Brooks 
(1962a, 1969), as well as information from new material from Brazil (Dr. Iraja Domiani 
Pinto, pers. comm.), reveals that there are 8 free subequal thoracomeres, placing this 
species within the Palaeocaridacea. 

Clarkecaris brasilicus (Clarke) 1920 
Fig. 25; Plate 10, figs. B-D 

\.*1920 Gampsonyx brasilicus. Clarke, p. 137, pi. 3, figs. 9, 10. 

1927 Uronectes brasilicus (Clarke). Chappuis, p. 605. 

1931 Uronectes braziliensis (Clarke). Van Straelen, p. 18. 

1946 Gampsonyx brasilicus Clarke. Mezzlira, p. 118, figs. 9, 10. 

1946 Uronectes brasilicus (Clarke). Mezzalira, p. 118. 

1948 Uronectes brasilicus (Clarke). Mezzalira, p. 250. 

1952 Clarkecaris brasilicus (Clarke). Mezzalira, p. 46, pi. 3. 

1954 Clarkecaris brasilicus (Clarke). Mezzalira, p. 168. 

1959 Gampsonyx brasilicus Clarke. Slewing, p. 100. 

V. 1962a Clarkecaris brasilicus (Clarke). Brooks, p. 231; Fig. 2b; pi. 5, fig. 2. 

1962b Clarkecaris brasilicus (Clarke). Brooks, p. 274. 

1969 Clarkecaris brasilicus (ClaTke). Brooks, p. R358, figs. 169-3, 174-2. 

1969 Clarkecaris brasilicus (Clarke). Schram, p. 221, table 1. 

1971 Clarkecaris brasilicus (Clarke). Mezzalira, p. 319, pi. 1, fig. 1, pi. 3, figs. 1-6. 

1977 Clarkecaris brasilicus (Clarke). Schram, p. 370. 

1978 Clarkecaris brasilicus (Clarke). Brito and de Quadros, p. 417, fig. 3. 

1979 Clarkecaris brasilicus (Clarke). Schram, p. 170. 
1981 Clarkecaris brasilicus (Clarke). Schram, p. 130. 

Lectotype. -NYSM 9738 (Plate 10, fig. D); near Guare'i, Sao Paulo, Brazil; Irati 
Formation, Permian. 


Other /oca//7/>5. — Innumerable localities for this species are known in the states of 
Panama and Sao Paulo, in Brazil. These are summarized in Mezzalira (1948, 1954) 
and Brito and de Quadros (1978). 

Z)6'5<:/'//7//o«.— Cephalon with well-marked groove (Plate 10, fig. C). Antennules 
large, with well-developed peduncles medially serrate and setose. Antennae with large, 
oval, setose scaphocerite. Flagella well developed. 

Thoracomeres short, subequal in length, anterior margins marked with row of 
papillae, pleura apparently rounded. 

Abdominal segments long (Plate 10, fig. D), sixth longer than any others. Pleura 
various: first rounded, second through fifth with styliform posteriorly directed processes. 
Uropodal protopod with lateral styliform processes; exopod long and thin (possibly a 
straight diaeresis); endopod diaphanous, long, and oval. Telson shorter than uropod 
rami, narrow, subtriangular, terminus developed as bifed process (Plate 10, fig. B). 

Remarks.— T\iQ description here is based on observations derived from study of 
the types (NYSM 9738, 9739), USNM 1 12766, and the published figure in Brito and 
de Quadros (1978). Considerable more information should become available, however, 
as the collections available to Dr. Damiani Pinto of the Instituto de Geosciencias, Porto 
Alegre, Brazil, are eventually studied and described. These should allow a definitive 
placement of C. brasilicus within the palaeocaridacean families. 

Suborder ANASPIDACEA Caiman, 1904 

Infraorder ANASPIDINEA Caiman, 1904 

Family ANASPIDIDAE Thompson, 1894 

Genus ANASPIDITES Brooks, 1962a 

Diagnosis. — 'R.osXnxm. broad. Thoracomeres relatively short compared to anterior 
pleomeres. Telson subtriangular, distally pointed. 

Type species.— Anaspides antiquus Chilton, 1929. 

Remarks. — ThQ initial observations on this taxon (Chilton 1929, Brooks 1962a) 
were largely based on one incompletely preserved specimen. A search in 1980 of the 
reserve collections of the Australian Museum uncovered one additional specimen (F 
25226), which preserves the abdomen and parts of the tailfan. In addition, two spec- 
imens were found in the British Museum (Natural History). One of these (In 46114) 
is the finest example of this species known, revealing considerable details about ap- 
pendage anatomy. The other (In 46056) is of the uropodal exopods. Consequently, a 
redescription of the species and new reconstruction are presented here. 

Anaspidites antiquus (Chilton) 1929 
Fig. 26; Plate 10, figs. E&F 

v.* 1929 Anaspides? antiquus Chilton, p. 366, pi. 30. 

1962a Anaspidites antiquus (Chilton). Brooks, p. 234; pi. 5, fig. 1; figs. 1 & 2c. 

1962b Anaspidites. species Brooks. Brooks, pp. 267, 274. 

1969 Anaspidites antiquus (Chilton). Brooks, p. R356, figs. 169-4, 

J 982 Anaspidites antiquus (Chilton). Schram, p. 122. 

Diagnosis. — Since there is but one species known, the diagnosis of the species is 
the same as that of the genus. 

Holotype. — \JS 7903. Brookvale Brick Quarry, New South Wales. Hawksbury 
Sandstone, Triassic. 

Description.— Cephalon with broad rostrum, prominent cervical groove. Anten- 
nular peduncles large, with 3 subequal segments, flagella well developed [but length 
indeterminate because of lack of preservation]. Antennal peduncles with 4 (?) segments 
(US 7903). Mandibles large, massive. 

Thoracomeres somewhat shortened, almost one-half the length of anterior pleo- 
meres (US 7903, In 461 14). Pleura somewhat rounded (In 461 14). Thoracopods (Plate 
10, fig. F) with short coxae, bases, and ischia, long meri (In 461 14); beyond knee, long 



Plate 10 

Figure A. Brooksyncaris canadensis (Brooks) 1962, closeup of anterior end with cephalon (c) and first 4 
thoracomeres (numbered), xg.Q. 

Figures B-D. Clarkecaris brasilicus (Clark), 1920; B) paralectotype, NYSM 9739, showing telson with 
pointed tip (t) and styliform uropods (u), x7; C) USNM 1 12766, closeup of anterior end, note cephalon, 
size of thoracomeres (numbered) and anterior pleomeres (numbered), x 4.5; D) NYSM 9738, lectotype, with 
posterior thorax and abdomen, x 2.8. 

Figures E & F. Anaspidites antiquus (Chilton) 1929; E) F 25226, with abdomen and telson, x 1.8; F) In 
461 14, whole body, note annulate pleopods (arrows), x 1.6. 



Figure 26. Reconstruction of Anaspidites antiquus, scale 5 mm. 

carpi and propodi (US 7903, In 46114), dactyli incompletely preserved (In 46114). 
Neither thoracic epipodites nor exopods preserved. 

Pleomeres undecorated. Sixth pleomere length twice that of any anterior to it. 
Pleopods long, uniramous, annulate (In 46114, Plate 10, fig. F), protopods well de- 
veloped. Telson long, subtriangular, distally pointed (F 25226, Plate 10, fig. E) (perhaps 
some faint indication that terminus possibly flanked by set of small furcae). Uropodal 
protopod simple, well developed (In 46056, In 461 14). Exopod blade-like (F 25226, 
In 461 14), reinforced with thick struts along lateral and medial margins (In 46056). 

Remarks. — brooks (1962^) interpreted a 2-segment protopod on the antennae 
(with only the distal segment visible), a straight-edged scaphocerite, and the 3 most 
proximal joints of the flagellum as peduncular. I found no evidence for an antennal 
scale on either US 7903 or In 461 14. Brooks also felt that the thoracopods were widely 
spaced, on opposite ends of well-developed thoracic stemites. Close examination of 
US 7903 indicates that the supposed foramina of the thoracopods are more likely 
preservational anomalies of the cuticular wrinkles or possibly ridges on the anterior 
thoracomeres. Finally, the pleopod that Brooks noted is in fact part of a posterior 

Although the general mode of preservation of these fossils obscures much of the 
detail, enough can be discerned to be reasonably certain A. antiquus is an anaspid. The 
body is large and well developed, but the first thoracomere is fused into the cephalon, 
and the rami of the pleopods are clearly uniramous and annulate. However, the narrow 
thoracic somites and styliform telson clearly separate this Triassic species from the 
living forms found today in Tasmania. Unfortunately, the diagnostic features of the 
mouthparts are not visible on any of the available material of Anaspidites, and as a 
result exact assurance as to family affinities within the Anaspidacea must remain un- 


This research was carried out under NSF grant GB 79-03602, which allowed me 
to examine specimens in museums in Australia, Europe, and North America, and to 
study and collect living syncarids in the wilds of Tasmania. The following individuals 
were of direct help on various aspects of this study: Drs. H.-E. Griiner, Museum fur 
Naturkunde, Humboldt Universitat, Berlin; S. Morris, British Museum (Natural His- 
tory); S. Secretan, Institut de Paleontologie, Paris; G. Pacaud, Museum d'Histoire 
Naturelle, Autun; R. Prokop, Narodni Museum, Prague; A. Richardson and R. Swain, 
University of Tasmania, Hobart; H. K. Schminke, Universitat Oldenburg; G. Ubaughs, 
Laboratoire de Paleontologie Animale, Universite de Liege; H. van Amerom, Rijks 


Geologische Dienst, Heerlen; and R. Wilson, Institute of Geological Sciences, Edin- 
burgh. Photographic and technical assistance was rendered by Messrs. B. R. Burnett, 
R. M. Chandler, and T. A. Demere, San Diego Museum; and the reconstructions were 
drawn by Mr. M. J. Emerson. 

Literature Cited 

Berry, C.T. 1939. A summary of the fossil Crus- 
tacea of the order Stomatopoda and a descrip- 
tion of a new species from Angola. American 
Midland Naturalist 21:461-471. 

Boy, J. A. 1 972. Palokologischer Vergleisch zweier 
beriihmter Fossillagenstatten des deutschen 
Rotliegenden (Unterperm, Saar-Nahe-Gebiet). 
Notizenblatt des hessischen Landesamtes flir 
Bodenforschung 100:46-59. 

Brito, I. M., and L. P. de Quadros. 1978. Ocor- 
rencia inedita de Clarkecahs brazilicus no Per- 
miano do Estado de Parana. Anais Acadamia 
Brasileira Ciencias 50:417-421. 

Brocchi, P. 1880. Note sur un Crustace fossile 
recueilli dans les schistes d'Autun. Bulletin de 
la Societe Geologique de France (3)8:5-1 0, pi. 1 . 

Bronn, H. G. 1848. IJebeT Gampsonyx fimhriatus 
Jordan. Neues Jahrbuch fiir Minerologie, Geo- 
logie, und Palaontologie 1848:125-126. 

. 1850. Veher Gampsonyx fimhriatus Jor- 
dan aus der Steinkohlenformation von Saar- 
briicken und vom Murg-Thal. Neues Jahrbuch 
fiir Minerologie, Geologic, und Palaontologie 

Brooks, H. K. 1962a. On the fossil Anaspidacea, 
with a revision of the classification of the Syn- 
carida. Crustaceana 4:229-242. 

. 1962ft. The Paleozoic Eumalacostraca of 

North America. Bulletins of American Paleon- 
tology 44:163-338. 
-. 1969. Syncarida, pp. 345-359 in R. C. 

Stellung im System. Zoologisches Jahrbuch 40: 

1927. Anaspidacea, pp. 593-606 m W. 

Moore (ed.). Treatise on Invertebrate Paleon- 
tology, Part R, Arthropoda 4, Vol. 1 . Geolog- 
ical Society of America and University of Kan- 
sas, Lawrence. 

Burmeister, H. 1855. Ueber Gampsonychus fim- 
hriatus Jordan. Abhandlungen der naturfor- 
shenden Gesellschaft zu Halle 2:191-200, pi. 
10, figs. 12-14. 

Caiman, W. T. 1896. On the genus A naspides and 
its affinities with certain fossil Crustacea. 
Transactions of the Royal Society of Edinburgh 

. 1902. Uronectes and Anaspides, a reply 

to Prof Anton Fritsch. Zoologisches Anzieger 

. 1904. On the classification of the Mala- 

costraca. Annals and Magazine of Natural His- 
tory 13(7): 144-1 58. 

. 1911a. On Pleurocaris, a new crustacean 

from the English Coal Measures. Geological 
Magazine 8(5): 156-1 60. 

. 1911ft. On some Crustacea of the division 

of Syncarida from the English Coal Measures. 
Geological Magazine 8(5):488-495. 
-. 1932. Notes on Palaeocaris praecursor, a 

Kukenthal and T. Krumbach (eds.). Handbuch 
der Zoologie 111(1 ). de Gruyter, Berlin. 

Charig, A. J. 1982. Systematics in biology: a fun- 
damental companion of some major schools 
of thought, pp. 363-440 in K. A. Joysey and 
A. E. Friday (eds.). Problems of Phylogenetic 
Reconstruction. Academic Press, London. 

Chilton, C. 1929. Note on a fossil shrimp from 
Hawkesbury sandstones. Journal of the Royal 
Society of New South Wales 62:366-368. 

Clarke, J. M. 1920. Crustacea of the Permian of 
Sao Paulo, Brazil. Bulletin of the New York 
State Museum 219/220:135-137. 

Cockerell, T. D. A. 1916. The uropods oi' Acan- 
thotelson stimpsoni. Journal of the Washington 
Academy Science 6:234-236. 

Copeland, M.J. 1 957a. The Carboniferous genera 
Palaeocaris and Euproops in the Canadian 
Maritime Provinces. Journal of Paleontology 

. 1 957ft. The arthropod fauna of the Upper 

Carboniferous rocks of the Maritime Prov- 
inces. Memoirs of the Geological Survey of 
Canada 286:1-110. 

Dahl, E. 1983. Malacostracan phylogeny and evo- 
lution. Crustacean Issues 1:189-212. 

Eastman, C. R. (transl. and ed.). 1 900. Text-book 
of Paleontology (by K. A. Zittel). Macmillan & 
Co., London. 

Feistmantel, O. 1873. Ueber den Niirschaner 
Gasschiefer, dessen geologische Stellung und 
organische Einschliisse. Zeitschrift fiir deutsch- 
es geologisches Gesellschaft 25:593. 

Fritsch, A. 1 870. Ueber das Auffinden von neuen 
Thierresten aus der sogenannten Brettelkohle 
von Nysan bei Pilsen. Stizungsberichte der 
Koniglichen Bohmishen Gesellschaft der Wis- 
senschaften. Prag 1870:33-49. 

. 1875. Prirodopsis zivocisstva pro vyssi 

gymnasialni a realne skoly. Prague. 

. 1876. Fauna der Steinkohlenformation 

Bohmens. Archiv fiir naturwissenschaflliche 
Landesdurchforschung von Bohmen. Prag 2: 
1-15, 4 pis. 

. 1 90 1 . Fauna der Gaskohle und der Kalk- 

fossil crustacean of the division Syncarida. An- 
nals and Magazine of Natural History 10(10): 
Chappuis, P. A. 1915. Bathynella nutans und ihre 

stein der Permformation Bohmens 4:66-75. 

Goldenburg, F. 1877. Fauna Sarepontana Fossil- 
is. Die fossilen Thiere aus der Steinkohlenfor- 
mation von Saarbriicken 2:35-36. Saarbriick- 

Grobben, K. 1919. Ueber die Musculatur des 
Vorderkopfes der Stomatopoden der Crusta- 
ceen. Sitzungsberichte der Akademie der Wis- 
senschaften, Wien 101:237-274. 

Haack, W. 1927. Zur Kenntnis des Syncariden 
Uronectes (Gampsonyx) fimhriatus Jorden aus 


dem Rotliegenden. Preussisches geologisches 
Landesanstalt Jahrbuch 48:773-785. 

Hessler. R. R. 1983. A defense of the caridoid 
facies; wherein the early evolution of the Eu- 
malacostraca is discussed. Crustacean Issues 1 : 

Johnson, R. G., and E. S. Richardson. 1966. A 
remarkable Pennsylvanian fauna from the Ma- 
zon Creek area, Illinois. Journal of Geology 74: 

Jordan, H. 1847. Entdeckung fossiler Crustaceen 
im Saarbriicken'schen Steinkohlengebirge. 
Verhandlungen des naturhistorischen Vereines 
preussischen Rheinlande 4:89-92. 

Jordan, H., and H. von Meyer. 1854. Ueber die 
Crustaceen der Steinkohlenformation von 
Saarbriicken. Palaeontographica 4:1-8. 

Kent, L. S. 1982. Type and figured fossils in the 
Worthen collection at the Illinois State Geo- 
logical Survey. Circular of the Illinois State 
Geological Survey 524:1-65. 

Krejci, J. 1859. Eine neue Crustaces aus der boh- 
mischen Steinkohlenformation. Lotos 9:79-80. 

Kues, B. S., and K. H. Kitzke. 1981. A large as- 
semblage of a new eurypterid from the Red 
Tanks Member, Madera Fm. of New Mexico. 
Journal of Paleontology 55:709-729. 

Macmillan, D. L., G. Silvey, and I. S. Wilson. 1981. 
Coordination of the movements of appendages 
in the Tasmanian mountain shrimp Anaspides 
tasmaniae. Proceeding of the Royal Society of 
London (B)2 12:2 13-231. 

Malzahn, E. 1958. Ein neuer jungpalaozoischer 
Krebs aus dem niederrheinischen Zechstein. 
Deutsche geologishe Gesellschaft Zeitschrift. 

Matthews, S. C. 1973. Notes on open nomencla- 
ture and on synonymy lists. Paleontology 16: 

Meek, F. B., and A. H. Worthen. 1865. Notice of 
some new types of organic remains from the 
Coal Measures of Illinois. Proceedings of the 
Academy of Science, Philadelphia 1865:46-50. 

Meek, F. G., and A. H. Worthen. 1866. Paleon- 
tology of Illinois. Descriptions of Invertebrates 
from the Carboniferous System. Geological 
Survey of lUinois 2:399-406. 

and . 1868a. Preliminary notice of 

a scorpion, a Eurypterusl, and other fossils from 
the Coal Measures of Illinois. American Jour- 
nal of Science (2)46:27-28. 

and . 1868Zj. Paleontology of Illi- 
nois. Geological Survey of Illinois 3:289-565. 

Mezzalira, S. 1946. Novos crustaceos Paleozo- 
icos— crustaceos do Permiano de Sao Paulo, 
Brasil. Revista Instituto Geografico e Geolo- 
gico, Sao Paulo 4:1 15-1 18. 

. 1948. Distribui^ao dos fosseis do Estado 

de Sao Paulo. Mineragaoe Metalurgia 13:249- 

. 1952. Clarkecaris, nova genero de crus- 
taceos Syncarida do Permiano. Boletim de So- 
ciedade Brasileira de Geologia 1:46-51. 

. 1954. Novas ocorrencias de crustaceos 

fosseis da Formagao Irati do Sul do Brasil, pp. 
165-170 in F. W. Lang (ed.). Paleontologia do 
Parana. Comissao de Comemoraq:6es do Cen- 
tennario do Parana, Curitiba. 

. 1971. Contribufao ao conhecimento da 

geologia de sub-superficie e da paleontologia 
da Formacao Irati, no Estado de Sao Paulo. 
Anais Academic Brasileira Ciencias 43:273- 

Noodt, W. 1963. Crustacea Syncarida de Chile 
central. Investigaciones Zoologicas Chilenas 10: 

. 1965. Naturliches System und Biogeo- 

graphie der Syncarida. Gewasser und Abwasser 

Packard, A. S. 1885. The Syncarida, a group of 
Carboniferous Crustacea. American Naturalist 

. 1886a. On the Syncarida, a hitherto un- 

described synthetic group of extinct malacos- 
tracous Crustacea. Memoirs of the National 
Academy of Science 3:123-128. 

. 1886Z?. On the Gampsonychidae, an un- 

described family of fossil schizopod Crustacea. 
Memoirs of the National Academy of Science 

. 1889. Paleontological Notes. Notes on 

Carboniferous arthropods from Illinois. Pro- 
ceedings of the Boston Society of Natural His- 
tory 24:21 1-213. 

Paucaud, G., W. D. I. Rolfe, F. R. Schram, S. Se- 
cretan, and D. Sotty. 1981. Quelques inver- 
tebres nouveaux du Stephanien de Montceau- 
les-Mines. Bulletin de la Societe d'Histoire Na- 
turelle d'Autun 97:37-43. 

Peach, B. N. 1908. Monograph on the higher 
Crustacea of the Carboniferous rocks of Scot- 
land. Memoirs of the Geological Survey of Great 
Britain, Paleontology 1908:1-82. 12 pis. 

. 1914. On some Carboniferous arthropods 

with description of a new genus of myriapod. 
Proceedings of the Royal Physical Society 19: 

Pruvost, P. 1912. Note sur un myiapode du ter- 
rain houiller du Nord. Annales de la Societe 
Geologique du Nord 41:65-69. 

. 1919. Introduction a I'etude du terrain 

houiller du Nord et du Pas-de-Calais. La faune 
continentale du terrain houiller du Nord de la 
France. Memoires pour Servir a I'Explication 
des Cartes Geologiques Detaillee de la France 

. 1922. Description d'un crustace syncaride 

nouveau de I'assise de Chokier a Woensdrecht. 
Annales de la Societe Scientifique de Bruxelles 

. 1930. La faune continentale du terrain 

houiller de la Belgique. Memoires du Musee 
Royal d'Histoire Naturelle de Belgique 44: 107- 
282, 14 pis. 

Roemer, F. 1856. Palaeo-Lethaea: II. Theil. Koh- 
len-Periode (Silur-Devon-, Kohlen-, und Zech- 
stein-Formation), pp. 672-675 in H. G. Bronn's 
Lethaea Geognostica. Bd.I. Stuttgart. 

Rolfe, W. D. I. 1961. A syncarid crustacean from 
the Keele beds of Warwickshire. Paleontology 

, F. R. Schram, G. Pacaud, D. Sotty, and S. 

Secretan. 1982. A remarkable Stephanian 
biota from Montceau-les-Mines, France. Jour- 
nal of Paleontology 56:426-428. 

Sayce, O. A. 1908. Description of a new remark- 


able crustacean with primitive malacostracan 
characters. Annals and Magazine of Natural 
History l(8):350-355. 

Schminke, H. K. 1975. Phylogenie und Verbrei- 
tungsgeschichte der Syncarida. Verhandlungs- 
bericht der deutschen zoologischen Gesell- 
schaft 1974:384-388. 

. 1978. Die phylogenetische Stellung der 

Stygocarididae — unter besonderer Beriicksich- 
tigung morphologischer Ahnlichkeiten mit 
Larvenformen der Eucarida. Zeitschrift fur 
zoologische Systematik und Evolutionsfor- 
schung 16:225-239. 
-. 1981. Adaptation of Bathynellacea to life 

in the interstitial. Internationale Revue der ge- 
samten Hydrobiologie 66:575-637. 

Schneider, J., H. Walter, and J. Wunderlich. 1 982. 
Zur Biostratinomie, Biofazies, und Stratigra- 
phie des Unterrotliegenden der Breitenbacher 
Mulde (Thiiringer Wald). Freiberger For- 
schungsheft 366:65-84. 

Schram, F. R. 1969a. Stratigraphic distribution 
of Paleozoic Eumalacostraca. Fieldiana: Ge- 
ology 12:213-234. 

. \969h. Insights into the evolution of the 

Syncarida. Abstracts and Program of the An- 
nual Meeting of the Geological Society of 
America 1969:12. 

. 1976a. Some notes on Pennsylvanian 

crustaceans of the Illinois basin. Fieldiana: Ge- 
ology 35:21-28. 

. 1976^. Crustaceans from the Pennsyl- 
vanian Linton vertebrate beds of Ohio. Pa- 
leontology 19:411-412. 

. 1977. Palaeozoogeography of Late Paleo- 
zoic and Triassic Malacostraca. Systematic Zo- 
ology 26:367-379. 

. 1979a. British Carboniferous Malacos- 
traca. Fieldiana: Geology 40:1-129. 

. \919b. The Mazon Creek biota in the con- 
text of a Carboniferous faunal continuum, pp. 
159-190 in M. H. Nitecki (ed.). Mazon Creek 
Fossils. Academic Press, New York. 

. 1981a. Late Paleozoic crustacean com- 
munities. Journal of Paleontology 55:126-137. 

. 1981^. Stalking wild mountain shrimp 

'round the world. Part I. Environment South- 
west 493:8-1 1. 

— . 1981c. On the classification of the Eu- 
malacostraca. Journal of Crustacean Biology 1 : 

— . 1982. The fossil record and evolution of 
Crustacea, pp. 93-147 in L. G. Abele (ed.). The 
Biology of Crustacea Vol. I. Academic Press, 
New York. 

— . 1983. Method and madness in phylogeny. 
Crustacean Issues 1:331-350. 

— . 1984. Relationships within eumalacostra- 
can Crustacea. Transactions of the San Diego 
Society of Natural History 20(16):301-312. 

— , and J. M. Schram. 1979. Some shrimp of 

the Madero Formation (Pennsylvanian) Man- 
zanita Mts., New Mexico. Journal of Paleon- 
tology 53:169-174. 
Schram, J. M., and F. R. Schram. 1974. Squillites 
spinosus from the Mississippian Heath Shale 
of central Montana. Journal of Paleontology 

Scott, H. W. 1938. A stomatopod from the Mis- 
sissippian of central Montana. Journal of Pa- 
leontology 12:508-510. 

Scudder, S.H. 1882. Archipolypoda, a subordinal 
type of spined myriapods from the Carbonif- 
erous formation. Memoirs of the Boston So- 
ciety of Natural History 3:143-182. 

. 1 890. New Carboniferous Myriapoda from 

Illinois. Memoirs of the Boston Society of Nat- 
ural History 4:417-442. 

Secretan, S. 1967. Proposition d'une nouvelle 
comprehension et d'une nouvelle subdivision 
des Archaeostraca. Annales de Paleontologie 

. 1972. A propos d'un nouvel exemplaire 

du Syncaride d'Autun: Nectotelson rochei 
Brocchi. Bulletin de la Societe d'Histoire Na- 
turelle d'Autun 64:9-12. 

. 1980a. Les arthropodes du Stephanien de 

Montceau-les-Mines. Bulletin Societe d'His- 
toire Naturelle d'Autun 94:23-35. 

. 1980/?. Comparison entre des crustaces a 

cephalon isole, a propos d'un beau materiel de 
syncarides du Paleozoique implications phy- 
logeniques. Geobios 13:411-433. 

Slewing, R. 1959. Syncarida. in H. G. Bronn, 
KJassen und Ordnungen des Tierreiches 

Smith, G. 1909. On the Anaspidacea, living and 
fossil. Quarterly Journal of Microscopical Sci- 
ence 53:489-578. 

Thompson, G. M. 1893. Notes on Tasmanian 
Crustacea, with descriptions of new species. 
Proceedings of the Royal Society of Tasmania 

. 1894. On a freshwater schizopod from 

Tasmania. Transactions of the Linnean Soci- 
ety, Zoology 6:285-303. 

Van Straelen, V. 1931. Crustacea Eumalacostraca 
(Crustaceis decapodis exclusis), pp. 1-98 /«W. 
Quenstedt (ed.). Fossilium Catalogus I: Ani- 
malia. 48. W. Junk, Berlin. 

Vandenberghe, A. 1960. Un arthropode du ter- 
rain houiller de la Loire Eilecticus pruvosti. 
Bulletin de la Societe Geologique des France 

Vanhoffen, E. 1916. Die Anomostraken. Sit- 
zungsberichte Gesellschaft naturforschender 
Freunde zu Berlin 1916:137-152. 

Watling, L. 1981. An alternative phylogeny of 
peracarid crustaceans. Journal of Crustacean 
Biology 1:201-210. 

. 1983. Peracaridan disunity and its bear- 
ing on eumalacostracan phylogeny with a re- 
definition of eumalacostracan superorders. 
Crustacean Issues 1:213-228. 

White, C. A. 1 884. The fossils of the Indiana rocks. 
Reports of the Indiana Department of Geology 
and Natural History, Paleontology 13:107-180. 

Wood, S. P. 1982. New basal Namurian (Upper 
Carboniferous) fishes and crustaceans found 
near Glasgow. Nature 297:574-577. 

Woodward, H. 1881. Contributions to the study 
of fossil Crustacea. Geological Magazine (2)8: 

. 1908. Some Coal Measure crustaceans 

with modem representatives. Geological Mag- 
azine (5)5:385-396. 


— . 1911. On a large form of Anthrapalaeman 
from the clay-ironstone nodules of the Middle 
Coal Measures, Sparth Bottoms, Rochdale. 
Geological Magazine (5)8:361-366. 

Zittel, K. A. 1885. Handbuch der Zoologie Band 
I, Abteilung 2, Lief 4. Oldenbourg, Munich and 


Volume 20 Number 14 pp. 247-276 20 November 1984 

The Late Wisconsinan Vertebrate Fauna from Deadman Cave, 
Southern Arizona ,^ 

Jim I. Mead 

Center for the Study of Early Man. Institute for Quaternary Studies, 
University of Maine, Orono, Maine 04469 


Edward L. Roth J^^J^^fO 

Department of Biology, Howard Payne University, '^'^iMi/Tv, 

Brownwood, Texas 76801 *Y 

Thomas R. Van Devender 

Arizona- Sonora Desert Museum, Route 9, Box 900, 
Tucson, Arizona 85743 

David W. Steadman 

Division of Birds, Smithsonian Institution, 
Washington, DC. 20560 

Abstract. We report a particularly rich assemblage of fossil vertebrates from a cave in southern 
Arizona. This fauna provides new data for reconstructing the inadequately known Late Pleistocene- 
Early Holocene biota of the Sonoran Desert and nearby mountains. The vertebrate fauna of Deadman 
Cave includes 5 amphibians, 25 reptiles (13 lizards and 12 snakes), 12 birds, and 22 mammals for a 
total of 64 species. Only one amphibian {Bufo woodhousei), three reptiles (Callisaurus draconoides, 
Phrynosoma modestum, Gyalopium canum), and one mammal {Microtus species) are locally extirpated, 
although all still occur in southern Arizona. An unidentified icterine bird may prove to be an extinct 
species. Extinct mammals include Euceratherium collinum, Equus species, and Nothrotheriops shasten- 
sis, all large herbivores. Other than the extinct animals, the fauna dating of the Late Pleistocene and 
Early Holocene is little different from that which is available in southern Arizona today. What appears 
to have changed is the mosaic of the plant and animal community. Distinctly boreal animals are lacking 
from the fauna. The climate during the time of deposition of the cave sediment appears to have been 
equable; certain animals now confined to deserts were able to live in more diverse woodland com- 


Deadman Cave is a medium-sized, limestone cave at 1400 m (4600 ft) elevation 
on the northeastern side of the Santa Catalina Mountains, Pima County, Arizona (Fig. 
1). The cave is located in the lower portion of the mountain range where various 
Paleozoic limestone formations are exposed and it appears to be formed in the Mis- 
sissippian Escabrosa Formation (Wallace 1955). 

The present vegetation of this highly dissected area is desert-grassland intermixed 
with the lower boundary of the oak woodland {Quercus species); juniper {Juniperus 
erythrocarpa and J. deppeana) is thinly scattered (Whittaker and Niering 1968). Desert- 
grassland elements such as agave (Agave parryi'), ocotillo {Fouqieria sp/endens), variable 
prickly pear (Opuntia phaeacantha), and assorted grasses occur on the limestone and 
conglomerate hillslopes. Tributaries between the hills and the areas of the lower hill- 
slopes are covered with velvet mesquite {Prosopis velutina), netleaf hackberry (Celt is 



Figure 1 . Map of southern Arizona with the locations of Deadman, Papago Springs, and Ventana caves. 
Contour lines (1650 m, 5500 ft) denote the positions of large mountain ranges, illustrating that southeastern 
Arizona is mountainous with great expanses of woodland and boreal habitats. The southwestern portion of 
the state contains predominately small desert mountains. 

reticulata), and catclaw {Acacia greggii). The area around Deadman Cave is presently 
the ecotone between the creosotebush desertscrub communities of the San Pedro Valley 
(715m elevation at the town of Mammoth) and the oak woodland above. Between 
Deadman Cave and the top of the Santa Catalina Mountains (2790 m elevation) there 
are Mexican pine-oak woodland, and ponderosa pine and mixed conifer forests. 

The approximately three-by-five meter entrance to Deadman Cave is a collapsed 
cavern ceiling (Fig. 2). The cave once contained an elaborate system of active speleo- 
thems. Travertine building is now very rare to absent, possibly because of the devel- 
opment of the present entrance and resultant dessication; rimstone pools rarely contain 
any water (William Peachey 1980, personal communication). 

During the late 1800s miners entered the cave to explore the numerous passages. 
A cabin was constructed across the entrance to the cave. Possibly at the same time, a 
shaft was begun in a back portion of the cave. This shaft penetrated a travertine surface 
layer and 2.4 m of cemented rubble containing bones, and provided access to a lower, 




Figure 2. Generalized cross section of Deadman Cave, southern Arizona. The present entrance appears 
to have been the opening during the Late Pleistocene. A shaft built by miners during the 1800s cuts through 
two travertine layers and provides access to a lower room and the bone deposit, sealed off from the rest of 
the cave since approximately 8000 B.P. 


sealed-off, small room. Here a second travertine layer covered a fine carbonate silt. At 
this point a trench was excavated horizontally through this loose sedimentary layer 
and then all mining operations ceased, leaving the exposed shaft and trench walls. 

William Peachey informed us of the exposed sediments in Deadman Cave. One 
of us (ELR) and W. Peachey, entered the cave in 1972 and excavated an approximately 
1.0 by 0.5 by 0.5 m layer of loose sediment from below the travertine layer capping 
the material exposed in the trench wall. All excavated sediments were screened through 
window mesh (2 mm) sieves. Most of the fossils were identified using the comparative 
collections at the University of Arizona, Tucson and the Division of Birds, Smithsonian 
Institution, Washington, D.C. 


The faunal assemblage contains three extinct species: the Shrub ox {Euceratherium 
collinum), the Shasta ground sloth {Nothrotheriops shastensis), and the Horse {Equus 
species). The remainder of the fauna can be found today living in various habitats in 
southern Arizona. Euceratherium, Nothrotheriops, and Equus apparently were extinct 
along with many other large mammals by approximately 1 1 000 to 10 200 B.P. (years 
before present; Martin 1967; Mosimann and Martin 1975; Haynes 1968; Meltzer and 
Mead 1983). The youngest known radiocarbon date on dung of A^. shastensis is about 
10 500 B.P. (Thompson et al. 1980). These dates suggest that at least some of the 
Deadman Cave faunal assemblage is of Late Wisconsinan age. 

A radiocarbon date of 6080 ± 250 B.P. (A-1617) was determined on 14.8 grams 
of endocarps of Cehis reticulata found directly associated with the fauna. Unfortunately 
there was insufficient CO2 available for a '^C correction. Cehis endocarps are notorious 
for containing very little carbon and for being easily contaminated by carbonates in 
percolating water. The sediments had been leached of all organics; some of the bones 
were encrusted with carbonates. For this reason we believe that the ''^C age of 6000 
B.P. is probably too young by at least 2000 years. The Deadman Cave faunal assemblage 
presented here most likely dates between 12 000 and 8000 B.P., grading across the Late 
Wisconsinan-Early Holocene boundary. Because of the uncertainties of the dating it is 
not possible to establish unequivocally whether all the reported taxa lived contem- 
poraneously in the local community. The thick travertine cap and the sealing off' of the 
lower room suggests that the deposit has not been contaminated with Middle or Late 
Holocene bones. 

The Fossil Deposit 

The fossiliferous layer is a pebbly silt (very pale brown, 7/3 10 YR dry, Munsell 
color) and shows no physical indication that the sediments were deposited by flowing 
water. The bones may have accumulated by a number of mechanisms. Bassariscus 
astutus (Ringtail) is a small carnivore that inhabits the cave today; small pockets of 
bones and seeds are presently developing as the scats of the Ringtail decay. Some of 
the fossil bone deposit in Deadman Cave may have been developed by the Ringtail as 
it was 2000 B.P. in Vulture Cave, Grand Canyon, Arizona (Mead and Van Devender 
1981). Spilogale putorius (Spotted skunk) is the most common carnivore in the fossil 
deposit and it too may have helped in the accumulation of animal remains. 

Owls {Otus species— Screech owl; Micrathene whitneyi—E\{ o-wX; and Asio otus— 
Long-eared owl) were also recovered from the Deadman Cave deposit. The Long-eared 
owl is known to prey upon Spotted skunks and is the only owl that inhabits deep 
recesses in caves, that was recovered from the deposit. 

The Ringtail and the Long-eared owl could very well account for the entire fossil 
deposit except for the larger mammal remains. The cave also may have been a den 
and a food cache for Felis concolor (Mountain lion) or other large carnivores, which 
could account for the occasional fossil remains of the Mountain lion. Horse, Ground 
sloth, and Shrub ox. 

The area of the bone deposit (Fig. 2) is just beyond view of the light coming in 


from the present and the presumed Late Wisconsinan-aged cave entrance. From here 
to the area of the fossil deposit, it is an easy passage for an owl or a mammalian 
carnivore, across an open, large cavern room. The predators may have had easy access 
to the lower room where the fossil bone deposit occurs, but the passage has since filled 
in with travertine, bones, and rock rubble. 

Thus, the Deadman Cave fossil deposit is probably time transgressive by possibly 
a few thousand years, and appears to be an in situ deposit from the predator accu- 
mulation(s) dating some 8000 to 12 000 years ago. 


The vertebrate fauna recovered from Deadman Cave (University of Arizona, Lab- 
oratory of Paleontology, UALP, locality 78121), is represented by 5 amphibians, 25 
reptiles (13 lizards and 12 snakes), 12 birds, and 22 mammals, for a total of 64 species 
(Table 1). 

The following is a systematic account of the fauna. After the skeletal element is 
the number of specimens, if more than one, and the UALP specimen number. "R" 
and "L" refer to right and left respectively. A brief discussion of identifying criteria 
and present-past distributions within the southwestern United States and northern 
Mexico is included. Our taxonomic sequences, distributional data, and descriptive 
osteological nomenclature is as follows: amphibians and reptiles— Stebbins (1966); 
birds— Baumel et al. (1979), Phillips et al. (1964); and mammals— Jones et al. (1982) 
for extant species and Kurten and Anderson (1980) for extinct species. Amphibians 
and reptiles were identified by TRVD and JIM, birds by DWS, and mammals by ELR 
and JIM. 

Class AMPHIBIA— Amphibians 

Order ANURA —Toads and frogs 

Family Pelobatidae— Spadefoot toads 

Scaphiopus species— Spadefoot toad 

Material — Kdidio-uXna. (11411). 

Scaphiopus cowc/z/- Couch's spadefoot toad 

Ma/ma/.- Vertebra (1 1409); R ilium (15312). 

Remarks. — Skeletal elements from adult individuals of Scaphiopus couchi and S. 
hammondi are distinctive. The shape of the ilium, including the Ala ossa ilei and the 
Margo dorsalis, identify the fossil specimen as belonging to a spadefoot toad. Scaphiopus 
couchi may be distinguished from 5. hammondi (Western spadefoot toad) by the fol- 
lowing characters: 1) the area from the spina pelvis anterior to the acetabulum is 
relatively flat on S. hammondi but is raised at the acetabulum on S. couchi; 2) the 
articular surface at the acetabulum is curved on S. hammondi, but straight on 5'. couchi; 
and 3) the shape of the Spina pelvis posterior is angular on S. hammondi, but curved 
on 5". couchi. S. couchi lives in a wide variety of desertscrub, grassland, and subtropical 
habitats including the Santa Catalina Mountains. 

Amphibian subfossil and fossil remains are very inadequately represented in Ar- 
izona (Van Devender and Mead 1978, Mead 1981). Besides Deadman Cave, S. couchi 
is known from an Early Holocene age wood rat midden in Arizona (Van Devender 
and Mead 1978), Late Wisconsinan and Holocene cave deposits in southwestern New 
Mexico (Van Devender and Worthington 1977, Holman 1970, Brattstrom 1964), and 
from Rancho la Brisca, Sonora, Mexico (Van Devender et al. in press). 

Scaphiopus cf. S. hammondi— WestQm spadefoot toad 
Material. — Vertebra (11410). 


Table 1 . Late Pleistocene, Holocene and present fauna from Deadman Cave and other localities in southern 
Arizona. Sequence and nomenclature is as follows (exceptions— see text): the amphibians and reptiles 
follow Stebbins (1966), the birds follow Phillips et al. (1964), and the mammals follow Jones et al. (1982) 
for the extant species and Kurten and Anderson (1980) for the extinct species. 1 = Van Devender and 
Mead (1978); Van Devender (1973); Mead et al. (1983). 2 = Haury (1950). 3 = Skinner (1942) and Rea 
(1980). * = occurs in category. ! = extinct species. S.C.Mt. = Santa Catalina Mountains. ? = Questionable 

Present fauna 

Wood rat middens' 

Ventana Cave 
< 10,000 B.P.2 


or nearby Southern Deadman < 10,000 > 10,000 Volcanic Conglom- 
valley Arizona Cave B.P. B.P. unit erate 



Scaphiopus couchi 
S. cf 5'. hammondi 
Bufo cf B. wood- 

B. punctatus 
Rana sp. 


Gopherus agassizi 
Coleonyx variegatus 
Sauromalus obesus 
Holbrookia maculata 
H. texana 

Callisaurus draconoides 
Crotaphytus collaris 

C. wislizeni 
Sceloporus cf. 5". 

S. cf. 5'. clarkii 
S. cf. 5". undulatus 
Uta stansburiana 
Urosaurus cf U. 

U. ornatus 

Phrynosoma douglassi 
P. modestum 
P. solare 
Cnemidophorus cf. 

C tigris 
Cnemidophorus sp. 
Heloderma suspectum 
Lichanura trivirgata 
Phyllorhynchus decur- 

Masticophis sp. 
Salvadora sp. 
Arizona elegans 
Pituophis melano- 

Lampropeltis getulus 
L. pyromelana 
Rhinocheilus lecontei 
Sonora semiamdata 
Chionactis occipitalis 
Gyalopium canum 
Trimorphodon biscu- 

Hysiglena torquata 
Crotalus atrox 
C. cerastes 
C. scutulatus 















Table 1. Continued. 

Present fauna Ventana Cave Spnngs 

— — — Wood rat middens' < 10,000 B.P.= * , ^ 

S.C.Mt. : Late 

or nearby Southern Deadman < 10,000 > 10,000 Volcanic Conglom- Pleisto- 

valley Arizona Cave B.P. B.P. unit erate cene 


Ibis-like * 

Colinus gambelii 
Colinus sp. 

Cyrtonyx montezumae 
Meleagris crassipes 
Zenaida cf. Z. 

Olus sp. 

Micrathene whitneyi 
Asio otus 
Colaptes auratus 
Turdus cf. T. migra- 

Catharus guttatus 
Icterinae (probably 

extinct species) 
Emberizinae » * * 


* * « 

if * * 

itC * 3tt 

* * * 

* * * 

* * * 

* m * 

« He * 

* * « 

* * * 

* * * 

Notiosorex crawfordi 
Myotis cf M. velifer 
M. cf M. evotis 
M. cf M. thysanodes 
cf Myotis 
Plecotus cf P. rafin- 

Antrozous pallidas 
Tadarida cf. T. bra- 

\Nothrotheriops shas- 

Sylvilagus auduboni 
Sylvilagus sp. 
L. californicus 
Lepus sp. 
Eutamias dorsalis 
Eutamias sp. 
Marmota flaviventris 
cf Ammospermophilus 
Spermophilus varie- 

S. tereticaudus 
S. lateralis 

Cynomys ludovicianus 
Thomomys bottae or 

T. cf T. bottae 
Perognathus cf P. 

P. cf P. flavus 
P. baileyi 
Perognathus sp. 
Dipodomys spectabilis 
D. cf D. deserti 
D. merriami 
Reithrodontomys mon- 



* * * 

* * * 


* * * 

« * * * * 

* * 

* * 

* * 

* * 

* * 

* * 

* * 

* * 

* * 

* * 

* * * 

* * * 

* * * 

* * * 

* * 

* * * 

Table 1. Continued. 


Present fauna 

Ventana Cave 
Wood rat middens' < 10,000 B.P.^ 





or nearby Southern Deadman < 10,000 > 10,000 Volcanic Conglom- Pleisto- 
valley Arizona Cave B.P. B.P. unit erate cene 

Reithrodontomys sp. 



Peromyscus manicu- 




P. boylii or trueP 



Peromyscus sp. 




Onychomys torridus 



O. leucogaster 


Sigmodon cf. S. ari- 





S. ochrognathus 


Neotoma albigula 




N. lepida 


N. mexicana or albi- 




Microtus sp. 



M. cf. M. mexicana 


Erethizon dorsatum 



Canis latrans 



C. lupus 


!C. dirus 

Vulpes macrotis 



Urocyon cinereoar- 




Ursus americanus 



Bassariscus astutus 




\B. sonoitensis 

Taxidea taxus 



Spilogale putorius 




Mephitis mephitis 



M. macroura 




Felis concolor 




\Panthera leo atrox 

\Equus tau 

\E. occidentalis 

\E. conversidens 

\Equus sp. 


! Tapirus sp. 

\Platygonus comp- 


Dicotyles (=Tay- 

assu) sp. 



\Camelops sp. 

Odocoileus sp. 




Cen'us sp. 

IStockoceros cf. S. 


\S. onusrosagris 

\Euceratherium col- 



Bison bison 

Bison sp. 





Remarks. — The shape of centrum and the size of the fossil vertebra was indistin- 
guishable with that of the Western spadefoot toad. S. hammondi is widely distributed 
in southeastern Arizona where it lives in desert, grassland, chaparral, woodland, and 
pine forest habitats (Lowe 1964). The fossil referred to as cf. S. hammondi represents 


the first fossil occurrence for the species in Arizona. Late Pleistocene and/or Early 
Holocene age occurrences outside Arizona for this species are known from Nevada 
(Brattstrom 1976) and New Mexico (Holman 1970). 

Family Bufonidae— Toads  

Bufo cf. B. woodhousei—V\l oodhoxxst's, toad 

Material. -Y^nthrdi (1 1408). 

Remarks.— Bufo woodehousei is a large toad in relation to the other species found 
today in Arizona, although distinctly smaller than B. alvarius (Colorado River toad). 
Bufo woodhousei can be identified by: 1) the size is relatively larger at all stages of 
growth than most other Bufo, 2) the neural arch is higher making the centrum more 
pronounced, and 3) the articular facets are larger. Woodhouse's toad occurs in eastern 
and central Arizona and in isolated populations in the Yuma area of southwestern 
Arizona. In southern Arizona B. w. australis is primarily a riparian species restricted 
to permanent or semi-permanent streams. Late Pleistocene or Early Holocene age 
remains of Bufo woodhousei have not been previously reported from Arizona. Outside 
Arizona, fossil remains of this toad are known from Nevada (Mead et al. 1982) and 
New Mexico (Holman 1970). 

Bufo punctatus—Ktd-spoUQd. toad 

Material. - Urostyle ( 1 5002). 

Remarks.— Bufo punctatus is a small toad with many easily identifiable skeletal 
elements. The paired anterior condyles of the urostyle are relatively broad, flat ovals 
as in B. punctatus. Juveniles of the larger species of Bufo, do not have as flattened an 
anterior end to the urostyle. This xeric-adapted toad occurs throughout most of the 
Southwest, living in habitats ranging from desertscrub to Mexican pine-oak woodland. 
Late Pleistocene remains of B. punctatus occur in three Arizona localities (Van De- 
vender and Mead 1978), New Mexico (Holman 1970, Van Devender and Worthington 
1977), and Rancho la Brisca, Sonora, Mexico (Van Devender et al. in press). 

Family Ranidae— Frogs 
Rana species— Frog 

Material — WnmcYViS (11412). 

Remarks. — T\\Q long, slender humerus is identifiable to Rana, but we were only 
able to identify the fossil to a small species. Rana pipiens (Leopard frog) and R. 
tarahumarae (Tarahumara frog) along with the introduced R. catesbeiana (Bull frog) 
occur in southern Arizona today. Only R. pipiens and the introduced species occur 
near Deadman Cave where they are restricted to permanent water habitats along streams. 
Rana species have been recovered as Quaternary fossils in California (Brattstrom 
1953<2, b, Hudson and Brattstrom 1977), Nevada (Brattstrom 1954), New Mexico 
(Holman 1970, Van Devender and Worthington 1977), and Rancho la Brisca, Sonora, 
Mexico (Van Devender et al. in press). 

Class REPTILIA- Reptiles 

Order SQU AM AT A— Lizards and Snakes 

Suborder Sauria — Lizards 

Family Iguanidae— Iguanid lizards 

Holbrookia maculata— Lesser earless lizard 

Material. — Dentary ( 1 1 394). 

Remarks.— The Lesser earless lizard is a small ground-dwelling lizard common in 
open habitats of desertscrub, desert grassland, and oak woodlands. It lives in the lower 
elevations of the Santa Catalina Mountains and occurs over most of eastern Arizona 
(Lowe 1964). The only previous Late Pleistocene and Early Holocene records of this 
lizard are from New Mexico (Van Devender and Worthington 1977). 


Holbrookia texana— Greater earless lizard 

Material. -R & L dentaries (2; 11401); R maxilla (1 1395). 

Remarks. — Teeth of//, texana are relatively taller on a deeper dentary than those 
of the smaller //. maculata. Dentaries of Holbrookia have a closed but not fused 
Meckel's canal. Today this lizard is found in open habitats on the south side of the 
Santa Catalina Mountains, but not near the cave at present. This insectivorous lizard 
lives at middle elevations in west central and southern Arizona, avoiding extreme 
desert lowlands (Stebbins 1966, Lowe 1964). The only other known Late Pleistocene 
or Early Holocene records of this lizard are from New Mexico (Van Devender and 
Worthington 1977). 

Callisaurus draconoides— Zebra-tailed lizard 

Material. — L dentaries (2; 1 1386). 

Remarks. — Teeth and dentaries of Callisaurus draconoides are similar to those of 
most medium-sized sceloporine lizards and to Holbrookia in particular, but they can 
be differentiated using an ontogenetic size series of specimens. Dentaries and teeth of 
Callisaurus are much larger and more robust than those of species of Holbrookia; 
osteologically Callisaurus is most similar to //. texana. The anterior one third of the 
Callisaurus dentary is very slender and has a more medial, internal, orientation to 
Meckel's canal, as compared to the more ventral orientation of either Holbrookia or 
Sceloporus. The Zebra-tailed lizard lives in regions of fairly open sandy or gravelly, 
low-elevation, desertscrub communities. The nearest population to Deadman Cave is 
in the low areas near Florence Junction and along the San Pedro River. This is the first 
Late Pleistocene-Early Holocene record of C draconoides. 

Crotaphytus col laris— Collared lizard 

Material.— R & L dentaries (6; 11390); R & L maxillae (8; 11391); pterygoid 
(11392); frontal (15003). 

/^^war/c^. — Specimens of C collaris and C wislizeni (Leopard lizard) can be sep- 
arated from most other iguanid lizards by their overall larger size, and the tendency 
for the teeth to be pointed and recurved, an adaptation for their carnivorous habits. 
The teeth of C collaris are relatively wider anteroposteriorly than those of C wislizeni, 
with the posterior teeth strongly tricuspid and the anterior teeth being more like blunt 
cones with a slight posterior curve. Both the pterygoid and the frontal are more rugose 
on C. collaris than they are on C. wislizeni. 

We use the name Crotaphytus collaris (sensu lato) and have not tried to separate 
C. collaris from C. insularis (Smith and Tanner 1972, Montanucci et al. 1975). Collared 
lizards can be found in all mountainous regions of southern Arizona and occasionally 
on open flat terrain (Lowe 1964). It presently lives near Deadman Cave. 

The Collared lizard is known from fossil sites in Arizona (Van Devender and Mead 
1978, Mead 1981, Cole and Mead 1981), Nevada (Brattstrom 1 954a, Mead et al. 1 982). 
New Mexico (Holman 1970, Gehlbach and Holman 1974, Van Devender and Worth- 
ington 1977). 

Sceloporus cf. 5. clarkii— Clark's spiny lizard 
Material. -"L dentaries (2; 1 1396); R & L maxillae (2; 1 1397). 

Sceloporus cf. S. magister— Desert spiny lizard 
Material. — R & L dentaries (2; 1 1398). 

Sceloporus clarkii or magister— Clark's or Desert spiny lizard 

Material. -R & L dentaries (10; 1 1400); R & L maxillae (9; 1 1399). 

Remarks. — Osteologically it is difficult to distinguish these two moderately large 


spiny lizards in their southeastern Arizona range. Sceloporus magister can have more 
robust dental characters. It usually inhabits the low deserts but will occur up into the 
desert-grassland. Sceloporus clarkii lives in woodlands in Arizona, but is a common 
inhabitant in the subtropical thomscrub in Sonora, Mexico. Both species ^re found 
today in the Santa Catalina Mountains. Other Late Pleistocene and/or Early Holocene 
records of Clark's spiny lizard are in New Mexico (Van Devender and Worthington 
1977) and Rancho la Brisca, Sonora, Mexico (Van Devender et al. in press). Sceloporus 
magister is fairly common in the fossil record, including Arizona (Van Devender and 
Mead 1978, Mead 1981), California (Brattstrom 1953a, b), and New Mexico (Van 
Devender and Worthington 1977). 

Sceloporus cf. S. undulatus—EasXem fence lizard 

Material. — R & L maxillae (3; 1 1387); R dentaries (2). 

Remarks.— These specimens are from a small species of Sceloporus similar to 
either S. undulatus or S. occidentalis (Western fence lizard). They can be distinguished 
from juvenile S. magister or 5". clarkii by their more slender, taller teeth. Maxillae and 
dentaries are less rugose in the Fence lizard, but are larger in all aspects than the S. 
graciosus (Sagebrush lizard). We are not convinced that 5". undulatus, S. occidentalis, 
or S. virgatus (Striped Plateau lizard) can be reliably separated satisfactorily on skeletal 

Sceloporus undulatus presently occurs near the cave, while 5. occidentalis and S. 
graciosus occur farther north and S. virgatus occurs in southeastemmost Arizona. For 
this reason the material may be referred to S*. undulatus. The Eastern fence lizard 
habitat in Arizona ranges from forested mountains down into the desert-grassland. 

Remains of S. undulatus are known from Late Pleistocene and Early Holocene 
deposits in Arizona (Van Devender and Mead 1978, Mead 1981, Cole and Mead 1981) 
and New Mexico (Holman 1970, Van Devender and Worthington 1977). 

Urosaurus ornatus— Tree lizard 

Material. — L dentary (1 1402). 

Remarks. — Urosaurus ornatus may be differentiated from most small iguanids 
including U. graciosus (Long-tailed Brush lizard) by its more slender teeth and the 
presence of a small fused area of the Meckel's canal. The Tree lizard in Arizona occurs 
in a wide variety of habitats from low, hot deserts up to open pine-oak woodlands. 
Late Pleistocene-Early Holocene remains of the Tree lizard have been found in south- 
western New Mexico (Van Devender and Worthington 1977). The Deadman Cave 
specimen is the first fossil record for the species in Arizona. 

Phrynosoma douglassi—Shon-homed lizard 
Material. -R & L dentaries (6; 1 1385); R & L maxillae (8; 1 1384). 

Phrynosoma modestum— Round-tailed homed lizard 
Material. -R dentary (1 1382); L maxilla (1 1383); parietal (3; 15004-15006). 

Phrynosoma solare— Regal homed lizard 

Material. -Parietal (11 381); angular (15007); squamosal (15008). 

/^^mar/c^. — Species of Phrynosoma can be differentiated from one another by most 
bones of the skull, especially those which bear horns (see Figs. 1-8 in Reeve 1952). 
The dentary, maxilla, and parietal are very mgose in P. modestum and are easily 
differentiated from the similar species, P. platyrhinos (Desert homed lizard), which 
lacks rugosity. Size, shape, and omateness will differentiate P. solare from other species 
(^fe also Reeve 1952). 

Phrynosoma douglassi presently occurs in the higher forests, woodlands, and grass- 


land habitats in eastern Arizona and the Santa CataHna Mountains, whereas, P. solare 
Hves in the Sonoran Desert valleys and bajadas, and adjacent desert-grasslands. Phry- 
nosoma modestum is a characteristic Chihuahuan Desert animal found in desertscrub 
and desert-grassland habitats. Today it occurs no further west than Sulphur Springs 
Valley, 95 km east of the San Pedro River Valley. This is the first fossil record for P. 
solare; Phrynosoma modestum is recorded from New Mexico (Van Devender and 
Worthington 1977); P. douglassi is recorded from New Mexico (Gehlbach and Holman 
1974, Van Devender and Worthington 1977) and Nevada (Mead et al. 1982). 

Family Teiidae— Teiid lizards 
Cnemidophorus species— Whiptail lizard 

Material. -I. dentary (1 1389); L maxilla (1 1388). 

/^6'mar/c.s'. — Neither the dentary nor the maxilla allowed for specific identification. 
Five species of Whiptail lizards occur in southern Arizona (C burti, C. exanguis, C. 
arizonae, C. inornatus, and C tigris). 

Family Helodermatidae— Beaded lizards 
Heloderma suspectum— Gila, monster 

Material. — Venehra (1 1393). 

Remarks. — Vertebrae of Heloderma can be separated from the only other large 
lizard of comparable size in Arizona, Sauromalus obesus (Chuckwalla), because they 
lack zygantra and zygosphenes and the dorsal half of the cotyle is oval rather than 
subsquare to orbicular. 

The Gila monster occurs in Arizona from the southern half of the state north into 
the extreme northwestern comer. Living primarily in the lowlands of the Sonoran 
Desert and portions of the Mohave Desert, the venomous Gila monster also occurs 
less commonly in desert-grasslands, and rarely in the oak woodlands. Heloderma sus- 
pectum is common along the lower portions of the Santa Catalina Mountains but 
probably does not occur today at Deadman Cave. 

It is not known whether the Gila monster occurred in Arizona, California, and 
Nevada during the Late Wisconsinan glacial or if it was a Holocene immigrant from 
the Sonoran Desert lowlands in the Lower Colorado River Valley around the head of 
the Gulf of California in Sonora, Mexico. Inadequately dated Late Pleistocene-Holocene 
remains occur at Vulture Cave, Arizona, and Gypsum Cave, Nevada (Mead and Phillips 
1981, Brattstrom 1954a). 

Suborder Serpentes— Snakes 

Family Colubridae— Colubrid snakes 

Masticophis species— Racer 

Material. — W^nebvat (1 1; 1 1404). 

Remarks.— The vertebrae of Masticophis are similar to those of Coluber (Racer) 
and Salvadora (Patch-nosed snake) {see the remarks under the latter species). Masti- 
cophis may be identified by the following: 1) the cotyle-condyle length (cl) is up to 6.5 
mm, occasionally to 8.2 mm, 2) the ratio of the cotyle-condyle length in relation to 
the neural arch width (NAW) is between 1.48 and 1.75, 3) the accessory process is 
long, pointed, and mostly oblique to anterior, and 4) the ratio of the distance between 
the prezygapophyses (PR-PR) and that distance between the prezygapophysis and the 
postzygapophysis (PR-PO) is between 0.87 and 1.00. This same ratio for Coluber is 
0.98 to 1.25 (Auffenberg 1963). 

Snakes of the genus Masticophis are large, active, diurnal predators. Within the 
genus, identification to species is difficult; the vertebrae of Coluber constrictor (Blue 
Racer) are similar as well. Masticophis flagellum (Coachwhip) and M. bilineatus (So- 
noran whipsnake) occur near Deadman Cave, while M. taeniatus (Striped whipsnake) 
occurs in the mountains to the north and northeast. Late Wisconsinan and Early 


Holocene remains of Masticophis species are known from New Mexico (Van Devender 
and Worthington 1977), Arizona, and California (Van Devender and Mead 1978), 
Nevada (Mead et al. 1982) and from the interglacial age Rancho la Brisca, Sonora, 
Mexico (Van Devender et al. in press). 

Salvador a species— Patch-nosed snake 

Ma/^r/a/.— Vertebrae (3; 1 1405). 

Remarks.— T\it vertebrae of Salvadora are similar to those of Coluber and Mas- 
ticophis. All generally have thin dorsal spines, a well-defined, thin haemal keel, and a 
tendency for epizygapophyseal spines. Salvadora is different from the latter two species 
in having a relatively smaller neural canal and a smaller condyle (Holman 1962). We 
do not know of any vertebral characters that unequivocally separate the two species 
within this genus. Snakes of the genus Salvadora are small ground dwellers. S. gra- 
hamiae (Mountain patch-nosed snake) occurs in the mountains of southeastern Arizona 
in oak woodlands and above, whereas S. hexalepis (Desert patch-nosed snake) is widely 
distributed in southern and western Arizona, living below the chaparral and woodland 
edge. Both species live in the Santa Catalina Mountains. The genus was recovered from 
the inter-glacial deposit at Rancho la Brisca, Sonora, Mexico (Van Devender et al. 
in press) and from a Late Wisconsinan-Early Holocene cave deposit in New Mexico 
(Van Devender and Worthington 1977). 

Arizona elegans— Glossy snake 

Material.— Ytnobvae {9; 11413). 

/^^marfo. — Characters used to identify the vertebrae of ^. elegans are: 1) the cl is 
up to 3.5 mm, 2) the ratio of cl and NAW is between 1.08 and 1.25, 3) the high neural 
arch, 4) the neural spine is high and moderately thin, 5) there is a long thin accessory 
process which is rounded and oblique to the anterior, and 6) the cotyle is oval to 
subround (Van Devender and Mead 1978). This medium-sized nocturnal snake lives 
in deserts and grasslands of most of the Southwest as well as in northern Mexico. This 
snake is known from Late Wisconsinan and Early Holocene remains in New Mexico 
(Van Devender and Worthington 1977) and Arizona (Van Devender and Mead 1978). 

Pituophis melanoleucus— Bull or Gopher snake 

Material. - Vertebrae (10; 11418). 

Remarks.— Criteria, for identification are discussed in Auffenberg (1963), but those 
used here are: 1) the cl is up to 7.5 mm, 2) the cl/NAW ratio between 1.07 and 1.17, 
3) the neural arch is very high with the neural spine being high and thick, 4) the 
zygosphene is moderately or strongly convex from the anterior, 5) the accessory pro- 
cesses are short, pointed or blade-like, and 6) the cotyle is round, relatively large, and 
only slightly oblique (Van Devender and Mead 1978). Pituophis melanoleucus is a 
widespread North American snake that lives in a wide variety of habitats in Arizona 
up to about 3000 m (9900 ft). Fossils of the species are found in Arizona (Van Devender 
et al. 1977, Van Devender and Mead 1978, Mead 1981, Cole and Mead 1981), Cali- 
fornia (Brattstrom 1953a), Nevada (Brattstrom 1958, 1976, Mead et al. 1982), and 
New Mexico (Van Devender and Worthington 1977). 

Lampropeltis getulus— Common king snake 

Material-WertehvaQ (7; 11416). 

Remarks. — ¥oT the species identification characters, see L. pyromelana. Fossils of 
L. getulus are known from Arizona (Van Devender et al. 1977, Van Devender and 
Mead 1978, Mead and Phillips 1981, Mead 1981), California (Brattstrom 1976, Van 
Devender and Worthington 1977). 


Lampropeltis pyromelana—Sonoran mountain kingsnake 

Material. — WeriehTae (8; 11417). 

Remarks.— The haemal keel and subcentral ridges are well developed in the king- 
snakes. Lampropeltis getulus is a large species and has a sharp neural spine with rel- 
atively blunt accessory processes. Lampropeltis pyromelana is a small species and has 
a thin, low neural spine and has short, pointed accessory processes. Both species occur 
in the Santa Catalina Mountains. L. getulus is common over most of North America, 
whereas L. pyromelana is found in montane habits in Nevada, Utah, and south into 
Mexico. Fossil remains have been reported from New Mexico (Van Devender and 
Worthington 1977) and Nevada (Mead et al. 1982). 

Rhinocheilus /^con/^/— Long-nosed snake 

Material. -MQnobraQ (22; 11419). 

Remarks.— A\\h.o\xg\\ vertebrae of i?. lecontei superficially resemble those o^ Lam- 
propeltis getulus, they are readily distinguished using the following criteria: 1 ) the cl is 
up to 3.0 mm, 2) the ratio of the cl and the NAW is between 1.07 and 1.21, 3) the 
neural spine is often flat-topped, 4) the zygosphene is often flat from the anterior, 5) 
the accessory process is blunt, lateral or dorsal from the anterior, 6) the cotyle is round 
and narrower than the zygosphene, and 7) the subcentral ridges are well-developed, 
but less so than in Lampropletis getulus (Auffenberg 1963, Hill 1971, and Van Devender 
and Mead 1 978). This medium-sized, nocturnal snake is widespread in desert, grassland, 
subtropical thomscrub habitats in the Southwestern U.S. and northern Mexico. The 
snake probably occurs near the cave today. Late Pleistocene-Holocene fossils occur in 
New Mexico (Van Devender and Worthington 1977), Arizona and California (Van 
Devender and Mead 1978, Mead 1981) and Nevada (Mead et al. 1982). 

Gyalopium ca«wm— Western hook-nosed snake 

Material. — Vertebra (11414). 

Remarks. — VtrXehrae of Gyalopium canum are small but very broad for their 
length. The haemel keel is poorly developed and the cotyle and condyle are relatively 
large compared to those of Sonora semiannulata (Ground snake) and Chionactis oc- 
cipitalis (Banded sand snake). 

The Western hook-nosed snake is a small snake that lives in the desertscrub and 
desert-grasslands from southeastern Arizona to Trans-Pecos, Texas and south into the 
Chihuahuan Desert of Mexico. Presently it occurs no closer to Deadman Cave than 
the Santa Rita Mountains, 80 km to the south. The only previous fossil record of the 
species (as Ficimia cana) was from New Mexico (Van Devender and Worthington 

Trimorphodon biscutatus— Lyre snake 

Material. -YeTXebrae (105; 1 1420). 

Remarks. — Chteha for the identification of T. biscutatus are as follows: 1) the cl 
is up to 4.5 mm, 2) the cl/NAW ratio is between 1.08 and 1.25, 3) the neural arch is 
flattened, 4) the neural canal is relatively small, 5) the zygosphene is relatively small. 
6) the accessory process is short and pointed, 7) the cotyle is oval to subround, strongly 
oblique, narrower than the zygosphene, and 8) the haemal keel is well-developed but 
low (Van Devender and Mead 1978). The Lyre snake is a medium-sized species that 
lives in desertscrub habitats in the Southwest and northern Mexico. It is found near 
Deadman Cave today. Fossils occur in New Mexico (Van Devender and Worthington 
1977), Arizona, and California (Van Devender and Mead 1978). 

Hypsiglena torquata—Nighl snake 
MateriaL-Yenebrae {\2- 11403, 11415). 


Remarks.— Criieria for identification are as follows: 1) the cl is between 1.65 and 
2.75 mm, 2) the cl/NAW ratio is 1 . 1 8 to 1 .3 1 , 3) the neural arch is moderately depressed 
from the posterior, 4) the neural spine is low, usually with the dorsal edge thickened 
and the anterior comer is bifurcate, 5) the accessory process is lateral from the anterior, 
and 6) the cotyle is relatively small (Van Devender and Mead 1978). The Night snake 
is widespread in desert, grassland, and woodland habitats in the Southwestern U.S. 
and northern Mexico. It occurs today near Deadman Cave. Fossils are known from 
New Mexico (Van Devender and Worthington 1977), Arizona and California (Van 
Devender and Mead 1978, Mead 1981), Nevada (Mead et al. 1982), and Rancho la 
Brisca, Sonora, Mexico (Van Devender et al. in press). 

Family Viperidae (= Crotalidae)— Pit vipers 
Crotalus a^rox— Western diamondback rattlesnake 

Material — YtnobraQ (5; 1 1406). 

Remarks.— Tht thoracic vertebrae of the Viperidae are distinct from those of the 
Colubridae and Boidae in that they have a long, pointed hypophysis. The fossil vertebrae 
are from a large rattlesnake resembling C. atrox; other rattlesnakes in Arizona generally 
do not attain its size, except for some C. molossus (Blacktailed rattlesnake). Both species 
occur near Deadman Cave. Crotalus atrox is a large desert species usually occuring in 
the lower valleys, whereas, C molossus is a woodland-dwelling species that is occa- 
sionally found in rocky habitats in more xeric desert mountain ranges. Late Pleistocene 
fossils of C. atrox have been reported from Gypsum Cave, Nevada (Brattstrom 1 954(2, 
b) and Conkling Cavern, Shelter, Fosbert (Brattstrom 1964) and Dry caves. New 
Mexico (Holman 1970). There are no unequivocal records of fossil C. atrox from 
Arizona; however, there is the interglacial record from Rancho la Brisca, Sonora, Mexico 
(Van Devender et al. in press). 

Crotalus scutulatus— Mohave rattlesnake 

Material. — Vertehme (3; 1 1407). 

Remarks.— The fossil vertebrae are from a medium-sized rattlesnake that is smaller 
than Crotalus atrox or C. molossus. The vertebrae of C. viridus cerberus (Arizona black 
rattlesnake), a common snake in the oak woodland, differ from those of C. scutulatus 
in their relative size of the hypophysis {see also Brattstrom \96Ab). The vertebrae of 
C. cerastes (Sidewinder), C lepidus (Rock rattlesnake), C. pricei (Twin spotted rattle- 
snake), and C. willardi (Ridgenosed rattlesnake) are smaller and differ in various mor- 
phological characters. The Mohave rattlesnake is a common desert-grassland and des- 
ertscrub snake in southern Arizona and near Deadman Cave today. Fossils of C 
scutulatus have not been reported previously. 

Class AVES- Birds 

Order GALLIFORMES- Gallinaceous birds 

Family Phasianidae— Pheasants, quails, etc. 

Colinus gambelii—GamheVs quail 

Material. -CormplelQ carpometacarpus (15313). 

Remarks.— This specimen is much smaller than the carpometacarpi of C ("'Or- 
eortyx"") pictus (Mountain quail) or Cyrtonyx montezumae (Harlequin quail), and is 
slightly smaller than that in C. {"'Callipepla'") squamata (Scaled quail). It differs from 
C. squamata and C. virginianus (Bobwhite) in having the Os metacarpale minus (meta- 
carpal III) more slender and more curved in caudal aspect, and in having a slightly 
smaller Processes extensorius. This is only the second fossil occurrence of C. gambelii, 
the other being from the Early Pleistocene (Irvingtonian) of Vallecito Creek, California 
(Howard 1 963). Brodkorb ( 1 964) listed C. gambelii questionably from Conkling Cavern 
and Shelter Cave, New Mexico. These assignments conflict, however, with the original 
references, as Howard and A. H. Miller (1933) reported ""Lophortyx sp. Quail" 


(sic) from these two sites. Gambel's quail lives today in the vicinity of Deadman Cave, 
but is approximately at its upper elevational limit. 

Colinus species— quail 

Material. — VroxxmaX and distal ends of humeri (15009). 

Remarks.— Among quail of the Southwest, these specimens are smaller than the 
humeri of Colinus {""Oreortyx") pictus or Cyrtonyx montezumae. We cannot, however, 
distinguish them from humeri of Colinus virginianus, C. {"" Lophortyx") gambelii, or 
C. C'Callipepla"") squamata, any of which could have occurred at Deadman Cave. 
Although species-level identifications are often very difficult, quail of the genus Colinus 
are common as Pleistocene fossils in southern North America, especially Florida, New 
Mexico, and California. 

Cyrtonyx montezumae— HarlQquin quail 

Material. — PTOximal end with partial shaft of radius (15010). 

Remarks.— This specimen is larger than the radius in all other southwestern quail 
except Colinus pictus. It is referable to Cyrtonyx by the less expanded articulating 
surface of the proximal end relative to the width of the shaft. The only other fossil 
occurrence of C montezumae is at San Josecito Cave, Nuevo Leon, Mexico (L. Miller 
1 943), also of Late Pleistocene age. Today the Harlequin quail occurs in grassy mountain 
woodlands of central and southeastern Arizona, thence ranging south well into Mexico. 
This species is very characteristic of evergreen oak grassland and is at its lower ele- 
vational limit near Deadman Cave today. With the historical reduction of grass and 
increase in brush at mid-elevations in Arizona mountains, this once common bird has 
decreased in abundance. 

Order COLUMBIFORMES- Pigeons and doves 

Family Columbidae— Pigeons and doves 

Zenaida cf Z. macrawra— Mourning dove 

Marma/. — Proximal end of carpometacarpus (1501 1). 

Remarks.— This fossil differs markedly in size from the carpometacarpi of all 
Arizona columbids except Zenaida macroura and Z. asiatica (White-winged dove). It 
is tentatively assigned to Z. macroura in being slightly smaller than all available spec- 
imens of Z. asiatica. This is the first Pleistocene record of Z. macroura in Arizona, 
although this species is a fairly common Late Pleistocene fossil elsewhere in North 
America. The Mourning dove is very widespread in Arizona, both geographically and 
altitudinally, and thus is of little paleoecological interest. 


Family Strigidae— Typical owls 
Otus species— Screech-owl 

Material.— Tv^o proximal ends and one distal end of humeri (3; 15012), proximal 
end of carpometacarpus (15013). 

Remarks. -These specimens all agree in size and morphology with Otus asio (Com- 
mon screech-owl), and are either larger or smaller than in all Arizonan owls outside of 
the genus Otus. The carpometacarpus and one proximal end of humerus are slightly 
larger than in O. Jlammeolus (Flammulated screech-owl), but the other elements re- 
semble both O. asio and O. jlammeolus. No skeleton was available for O. trichopsis 
(Spotted screech-owl), so identification beyond generic level is not possible. These three 
species of Otus in Arizona are largely separated from each other today by habitat and 
elevation, and it seems most likely that O. asio or O. trichopsis would have lived near 
Deadman Cave in the Late Pleistocene. Probably only O. asio occurs in the immediate 
vicinity of Deadman Cave today. Otus asio is a common Late Pleistocene fossil in 


North America, while O. flammeolus and O. thchopsis have only two and one Pleis- 
tocene records, respectively. This is the first fossil record of Otus in Arizona. 

Micrathene Whitney i— Elf o'wl * 

Material. — Proximal end of humerus (1501 4), distal end of tarsometatarsus (1501 5). 

Remarks.— ThQ Elf owl is readily separated from all other owls by its extremely 
small size. This is the first Pleistocene record for M. whitneyi. It occurs today in the 
region of Deadman Cave, nesting in holes in trees at any elevation "below the heavy 
pine forest" (Phillips et al. 1964). 

Asio o?W5— Long-eared owl 

Material — 'D\s\.3\ end of humerus (15016). 

Remarks. — This fossil agrees with the humerus of Asio otus versus A. flammeus 
(Short-eared owl) in having a distinctive knot-like ectepicondylar prominence (Pro- 
cesses supracondylaris dorsalis). Asio otus is a fairly common Late Pleistocene fossil 
in western North America, but this is the first such record in Arizona. The Long-eared 
owl is not unexpected at Deadman Cave, as it occurs today in Arizona in a variety of 
habitats, both as a nesting bird and a winter visitor. As mentioned above, A. otus 
probably was involved in the accumulation of small vertebrates in Deadman Cave. 

Order CAPRIMULGIFORMES- Goatsuckers, etc. 

Family Caprimulgidae— Night jars 

Genus and species indeterminate 

Ma?m<3/. — Carpometacarpus lacking distal end and much of metacarpal III (1 50 1 7). 

Remarks. — This carpometacarpus is distinguished from that of Chordeiles minor 
(Common nighthawk) and C. acutipennis (Lesser nighthawk) by its much smaller size, 
and from Caprimulgus vociferus (Whip-poor-will) by its slightly smaller overall size 
with a more slender metacarpal III. It resembles that of Phalaenoptilus nuttallii (Poor- 
will) very closely, but the shape of metacarpal I is somewhat more similar to that in 
Caprimulgus. In the absence of a comparative skeleton of Caprimulgus ridgwayi (Ridge- 
way's whip-poor-will), the only other caprimulgid living in Arizona, precise identifi- 
cation of this fossil is impossible. The Pleistocene record of caprimulgids is poorly 
known everywhere. This is the first fossil record for the family in Arizona. 

Order PICIFORMES- Woodpeckers, etc. 

Family Picidae— Woodpeckers 

Colaptes auratus—¥hc\<.tr 

Material. — Distal end of tarsometatarsus (1501 8). 

Remarks.— Among Arizonan woodpeckers, the tarsometatarsus of Colaptes au- 
ratus is similar in size only to that of Melanerpes {^" Asyndesmus"") lewis (Lewis' wood- 
pecker). The fossil agrees with C. auratus versus M. lewis in its larger, less deeply 
sculptured middle trochlea. The distal end of the tarsometatarsus in the "Red-shafted" 
flicker (C a. collaris) appears to be indistinguishable from that in the "Gilded" flicker 
(C a. mearnsi). Thus the fossil provides no evidence of paleohabitats near Deadman 
Cave. C. a. collaris is a bird of mountain woodland and forest, ranging upward from 
approximately 1220 m elevation, whereas C. a. mearnsi occurs in desertscrub, generally 
below 1 370 m elevation. Based on the remainder of the avifauna, one would guess that 
the Late Pleistocene flicker at Deadman Cave was C a. collaris, although both forms 
occur in the general region of the site today. Flickers are very common Late Pleistocene 
fossils, yet once again this is the first such record for Arizona. 


Order PASSERIFORMES- Perching birds 

Family Turdidae — Thrushes 
Turdus cf. T. migratorius— American robin 

Material. — Proximal end of humerus (15019). 

Remarks.— The humerus of Turdus migratorius can be told from that of most 
other North American turdids by its larger size. It can be recognized from that in 
Ixoreus naevius (Varied thrush) by its stouter Crus dorsalis fossae which, along with 
the deeper dorsal Fossa pneumotricipitalis, also distinguishes it from the humeri of 
mimids (thrashers, mockingbirds). The humerus of T. migratorius can be separated 
from that in the neotropical T. grayi (Clay-colored robin) reported from Stanton's 
Cave, Coconino County, Arizona (Rea and Hargrave, ms) by its lesser degree of pneu- 
maticity in both the dorsal and ventral Fossa pneumotricipitalis, the former also being 
larger in T. grayi. The fossil differs from the only available humerus of T. rufopalliatus 
(Rufous backed robin, a vagrant to Arizona today; resident in Sonora) in having a 
larger and more oblong (less circular) opening of the ventral Fossa pneumotricipitalis. 
Lacking additional specimens of T. rufopalliatus to confirm this character, no more 
than a tentative assignment of the fossil to T. migratorius is warranted. T. migratorius 
is a common Late Pleistocene species in much of North America, and has been reported 
in Arizona from Stanton's Cave (Rea and Hargrave, ms). The American robin is 
common in Arizona today, nesting throughout the state in wooded regions above 
approximately 1220 m elevation. 

Catharus guttatus-^HermiX thrush 

Material. — 'DisXaX end of humerus with most of shaft (19020). 

Remarks.— The humerus of Catharus guttatus is smaller than in mimids and in 
the following species of turdids: Hylocichla mustelina (Wood thrush), Ixoreus naevius, 
Myadestes townsendi (Townsend's solitaire), and all species of Turdus. It is larger and 
has a stouter Corpus humeri (shaft) than in T. ustulatus (Swainson's thrush). It is 
approximately equal in size to that of C. fuscescens (Veery), C. minimus (Gray-cheeked 
thrush), and the species of Sialia (bluebirds), but is told from these and all other Arizona 
turdids by having a relatively smaller Processus supracondylaris dorsalis, this being 
particularly evident in dorsal aspect, where P dorsalis is seen not to extend as far 
proximally in C guttatus as in other species. 

This is the first fossil record anywhere for C. guttatus. Brodkorb (1978) listed C. 
guttatus from the Late Pleistocene site of Carpinteria, California, citing A. H. Miller 
(\932b) as the authority. However, A. H. Miller (1932^^) clearly did not refer the 
specimen in question, a humerus, to any species. Miller listed the specimen as "HY- 
LOCICHLA? Thrush" (sic), stating that the fossil resembled Hylocichla {=Catharus) 
guttata in certain aspects, H. mustelina in others, and probably represented an extinct 
taxon of thrushes. The Hermit thrush is widespread in Arizona today, and is common 
in the Santa Catalina Mountains, nesting at high elevations and occurring elsewhere 
as a migrant or wintering bird. 

Family Fringillidae— Sparrows, finches, tanagers, blackbirds, warblers, etc. 

Subfamily Icterinae— Blackbirds, etc. 
Genus and species indeterminate 

Material. — Distal end of tarsometatarsus (15021). 

Remarks.— This specimen is larger than the tarsometatarsus in all non-icterine, 
nine-primaried oscines of the Southwest. Among southwestern icterines, it is smaller 
than in Sturnella magna (Eastern meadowlark), S. neglecta (Western meadowlark), 
and Cassadix mexicanus (Boat-tailed grackle), and larger than in Molothrus ater (Brown- 
headed cowbird) or any species of Icterus (orioles). Of the species that it approximates 
in size, the fossil may be distinguished: from Agelaius phoeniceus (Red-winged black- 


bird) and Xanthocephalus xanthocephalus (Yellow-headed blackbird) by its more dorso- 
plantar expansion of the middle and inner trochleae; from Euphagus cyanocephalus 
(Brewer's blackbird) and E. carolinus (Rusty blackbird) by its larger intertrochlear 
spaces, the more dorso-plantar expansion of the inner trochlea, and the more proximo- 
distally expanded outer trochlea; and from Molothrus aeneus (Bronzed coWbird) by its 
slightly wider outer intertrochlear space and slightly more laterally compressed inner 
trochlea. Overall, this specimen seems to be more similar to the tarsometatarsus of 
Molothrus aeneus than to any other living icterine, but is not similar enough to be 
referred confidently to that species. 

The fossil icterine from Deadman Cave may represent an extinct taxon. Four 
species of extinct icterines have been described from rostra and mandibles from Late 
Pleistocene sites in North America. These are Pandanaris convexa (A. H. Miller 1947), 
and Euphagus magnirostris (A. H. Miller 1929), both from Rancho La Brea, California; 
Pandanaris floridana (Brodkorb 1957) from Reddick and Haile XIB, Florida; and 
Pyeloramphus molothroides (A. H. Miller 1932a) from Shelter Cave, New Mexico. 
Referred post-cranial elements have been reported for Pandanaris floridana and Eu- 
phagus magnirostris, but we have not examined this material. All Late Pleistocene 
icterines are in need of re-study (Steadman and Martin, in press), and pending such 
work the specimen from Deadman Cave is best left unidentified. Nevertheless, it may 
represent a new faunal element for Arizona. 

Subfamily Emberizinae— "New World" sparrows, finches, etc. 
Genus and species indeterminate 

Afafm^/.— Tarsometatarsus lacking proximal end (15022). 

i^^mar/c^.— Postcranial emberizine fossils are often very difficult or impossible to 
identify to genus or species. The present specimen is smaller than the tarsometatarsi 
of any icterine (blackbirds) or North American thraupine (tanagers), and is smaller than 
in most parulines (New World warblers). It differs from the tarsometatarsi of vireonids 
(vireos) in its more slender middle trochlea and broader inner trochlea, this last char- 
acter also distinguishing it from the tarsometatarsi of parulines. Within the emberizines, 
no readily apparent patterns of tarsometatarsal variation are discernible. When com- 
pared to all species of North American emberizines, the fossil was found to be indis- 
tinguishable, both in size and quality, from the following species of medium-sized 
sparrows: Ammodramus sandwichensis (Savannah sparrow), Melospiza lincolnii (Lin- 
coln's sparrow), M. georgiana (Swamp sparrow), Junco hyemalis (Dark-eyed junco), 
and J. phaeonotus (Mexican junco). Geographical and sexual variation combine to 
render the tarsometatarsus of these five species inseparable in many instances. Certain 
individuals of each species appear to be distinct, but no consistent variation is seen. 
Each of these species occurs today in southern Arizona, although in different habitats 
and in very different frequencies. 

Class MAMMALIA— Mammals 
Order INSECTIVORA-Insectivores 

Family Soricidae— Shrews 
Notiosorex crawfordi—DQseri shrew 

Material. -L mandible (3; 15023); R mandible (2; 15024). 

Remarks. — The shape of the mandibles and the presence of three unicusps on each 
jaw were the identifying characters. Notiosorex crawfordi occurs fairly commonly as 
fossils in Arizona (Mead and Phillips 1981, Mead et al. 1 983), New Mexico, and Texas 
(Harris 1 977). Presently the Desert shrew occupies a wide variety of ecological situations 
from semi-desertscrub to woodland (Armstrong and Jones 1972). It is not known to 
occur at present in the Santa Catalina Mountains, but it does live nearby (Cockrum 



Family Vespertilionidae— Vesperlilionid bats 

cf. Mvo//5— Mouse-eared bat 

Material. -h mandible with M. (15025); R mandible (3; 15027). 

Remarks.— T\iQSQ specimens, clearly a vespertilionid based on the shape of the 
jaw, could not be identified unequivocally to genus because of fragmentation and/or 
for loss of teeth. Skinner ( 1 942) reported Myotis cf. M. velifer (Cave myotis), M. cf. 
M. thysanodes (Fringed myotis), and M. cf. M. evoltis (Long-eared myotis) from Papago 
Springs Cave, Arizona. 

Antrozous pallidus— Pallid bat 

Material. -L femur (15025). 

Remarks.— The greatest length of the fossil femur is 19.0 mm and the width of 
the proximal end is 3.0 mm (lesser trochanter to greater trochanter). A blade-like third 
trochanter is present. The Deadman Cave specimen was compared to Myotis thysan- 
odes, M. californicus (California myotis), M. velifer, Plecotus townsendii (Townsend's 
big-eared bat), Lasiurus borealis (Red bat), L. cinereus (Hoary bat), Macrotus water- 
housei (Leaf-nosed bat), Tadarida brasiliensis (Brazilian free-tailed bat), T. femorosacca 
(Pocketed free-tailed bat), Mormoops megalophylla (Ghost-faced bat), Antrozous pal- 
lidus, and Eptesicus fuscus (Big brown bat). Only the last two species were similar to 
the fossil in having a femur of total length averaging near 19.0 mm and a proximal 
width of 2.8 to 3.0 mm, along with the lesser trochanter as pronounced as the greater 
trochanter; but of these two, only A. pallidus had the third trochanter. The other species 
of bats lacked two or all of the criteria used to differentiate the fossil specimen. 

The Pallid bat occurs throughout Arizona and can be found near Deadman Cave 
today (Cockrum 1960, Barbour and Davis 1969). Bats of the genus Antrozous have 
been recovered as fossils in a wood rat midden in the Sonoran Desert (Mead et al. 
1983) and from Papago Springs Cave (Skinner 1942). 

Order EDENTATA -Edentates 

Family Megatheriidae— Megathere ground sloths 

Nothrotheriops shastensis —Shasta, ground sloth 

Material. — Molar (15314). 

Remarks.— Greg McDonald (Royal Ontario Museum, 1982, personal communi- 
cation) confirmed this identification of A^. shastensis and indicated that because the 
small molar contained a high percentage of hollow pulp cavity and lacked wear stria- 
tions, it must have been from a fetal or new bom sloth. Remains of the extinct Shasta 
ground sloth are very common in the Southwest, especially in Arizona (Long and Martin 
1974, Thompson et al. 1980). A typographical error in Lindsay and Tessman (1974) 
has the sloth incorrectly located in Stanton's Cave, Grand Canyon. 

Order LAGOMORPHA-Lagomorphs 

Family Leporidae— Hares and rabbits 

Sylvilagus species— Cottontail 

Material — 'L mandible; R mandibles (2); maxilla. 

Remarks. — VosXcramal remains of leporids were the second most common ele- 
ments in the fossil deposit. The mandibles and the maxilla are not identified to species 
at this time because a more detailed study of all Late Pleistocene leporid remains of 
Arizona is in order and will be appearing in the near future (JIM). The genus is recovered 
from a number of Late Pleistocene localities in Arizona (Lindsay and Tessman 1974, 
Mead et al. 1983) and New Mexico (Harris 1977). 


Lepus species— Jackrabbit 

Material — l. mandibles (6); R mandibles (4); L maxillae (2); R maxillae (2); pre- 
maxilla; isolated molars (4); L femur proximal half. 

Remarks.— See the remarks under Sylvilagus. Lepus alleni (Antelope jackrabbit) 
and L. callotis (White-sided jackrabbit) both presently occur in Arizona, but not near 
Deadman Cave. L. californicus occurs near Deadman Cave today. Lepus californicus 
(Black-tailed jackrabbit) was recovered from Papago Springs Cave (Skinner 1 942) and 
other Arizona localities (Lindsay and Tessman 1974, Mead et al. 1983). 

Order RODENTI A— Rodents 

Family Sciuridae— Squirrels 

Spermophilus variegatus—Rock squirrel 

Material. — L mandible with M,_2 (2; 15028); R mandible (15029); L maxilla with 

Remarks.— Spermophilus variegatus can be differentiated from other species of 
ground squirrels by its larger size and the tendency of the skeleton to be slightly more 
rugose. The only other squirrel of similar size is Sciurus aberti (Abert's squirrel). The 
P'* is relatively larger in Spermophilus variegatus as compared to that in Sciurus. The 
shape and medial inflection of the angle on the mandible is greater on S. variegatus. 
The Rock squirrel is a common ground squirrel and the largest within its distribution. 
It prefers rocky regions and is found throughout the Southwest, including the vicinity 
of Deadman Cave. Fossil remains of the Rock squirrel are not common in Late Pleis- 
tocene localities in the Southwest (Harris 1977, Mead 1981, Kurten and Anderson 
1980), although Skinner (1942) identified three mandibular rami of S. (^Citellus) 
variegatus from Papago Springs Cave. 

Family Geomyidae— Pocket gophers 
Thomomys cf. T. bottae— Bonne's pocket gopher 

Material. -L& R maxillae (3; 1 5036); isolated teeth (9; 1 5037); R humerus (1 5038). 

Remarks.— The upper incisors were lacking any conspicuous longitudinal groove, 
and the maxillary and isolated cheek teeth all were the lobbed, simple hypsodont molars 
and premolars of Thomomys. We follow Thaeler (1968) in using the designation T. 
bottae, which is the T. umbrinus of Hall (1981). The former species is common today 
in the Santa Catalina Mountains, and therefore, the reason for our identification of the 
fossils. The genus is a common fossil recovered in the Southwest (Mawby 1 967, Lindsay 
and Tessman 1974, Harris 1977). 

Family Heteromyidae— Pocket mice and Kangaroo rats 
Perognathus cf. P. flavus SiWay pocket mouse 

Material. -K maxillae (3; 15032); L maxillae (2; 15033); R mandibles (3; 15034); 
L mandibles (4; 15035). 

Remarks.— The pocket mouse specimens from Deadman Cave compare well with 
P. flavus, although two other indistinguishable, small pocket mice, P. parvus (Great 
Basin pocket mouse) and P. flavescens (Plains pocket mouse) could also be in the 
assemblage. Our tentative identification is based on the present geographic distributions. 
Complete skulls are needed for unequivocal identification. Perognathus flavus lives 
near Deadman Cave region. The only other Late Pleistocene occurrence of this mouse 
is from Isleta Cave, New Mexico (Harris and Findley 1964). 

Dipodomys 5/?£'c/<3Z>///5— Banner- tailed kangaroo rat 

Material. — Bacculum ( 1 503 1 ). 

Remarks.— The shape of the bacculum o^ Dipodomys spectabilis is distinct from 
that of all other species. Dipodomys spectabilis is a large kangaroo rat that inhabits the 


desert-grasslands of southeastern Arizona, including the valleys below the Santa Cat- 
alina Mountains (Cockrum 1960, Hall 1981). Harris (1977) has reported fossils of D. 
spectabilis from southern New Mexico. 

Family Cricetidae— New World Rats and Mice 
Reithrodontomys mo nt anus— Plains harvest mouse 

Material. -L mandible (15041); R mandibles (3; 15042). 

Remarks.— The mandibles and teeth of the harvest mice from Deadman Cave 
compare favorably with those of R. montanus. The other harvest mice in Arizona, R. 
megalotis (Western harvest mouse) and R. fulvescens (Fulvous harvest mouse), are both 
larger than R. montanus. The occlusal pattern of the molar of R. fulvescens is an "S" 
configuration as opposed to a "C" in R. montanus (Hooper 1952). 

Reithrodontomys montanus occurs today in the grasslands of southeastern Arizona 
but not in the Santa Catalina Mountains. The other two species occur in a wider variety 
of communities (Cockrum 1960). Fossil occurrences are discussed in Kurten and An- 
derson (1980). 

Peromyscus species— Deer mouse 

Materia!.— h mandibles (4); R mandibles (4). 

Remarks. — Fragmenls of Peromyscus can be confused with those of Reithrodon- 
tomys. The following characters will separate the two genera: 1) the M3 is relatively 
larger on Peromyscus, 2) the articular condyle of the mandible extends more posteriorly 
than does the angle, on Peromyscus, and 3) the angle of the mandible has a less medial 
inflection on Peromyscus. We were unable to identify these specimens to species. Eight 
species of Peromyscus occur in southern Arizona, thus species level identification of 
fossils is extremely difficult if not impossible. Fossils of the genus have been recovered 
from all over the Southwest (Harris 1977, Kurten and Anderson 1980, Mead et al. 

Sigmodon species— Cotton rat 

Material. -K mandible with M,_3 (15039); R maxilla with M'-- (15040). 

Remarks.— The occlusal pattern on all cheek teeth are distinct in Sigmodon. We 
are not able to identify our specimens to species. Four species of cotton rat now inhabit 
southern Arizona (Hall 1981). Sigmodon hispidus (Hispid cotton rat) occurs on the 
western and eastern borders of southern Arizona while Sigmodon arizonae (Arizona 
cotton rat) lives in the area of the Santa Catalina Mountains. Both 5. /w/v/'v^^f^T (Tawny- 
bellied cotton rat) and S. ochrognathus (Yellow-nosed cotton rat) occur south and east 
of Deadman Cave (Baker and Shump 1978a, b. Hall 1981). S. ochrognathus was 
recovered from a wood rat midden near the Santa Catalina Mountains (Mead et al. 
1983). The genus has a rich fossil record throughout the Southwest (Lindsay and 
Tessman 1974, Harris 1977). 

Neotoma a/Z7/gw/a— White-throated wood rat 

Material. — I. mandibles (3); R mandibles (4); L maxillae (3); R maxillae (7); LM, 
(7); RM, (8); LM" (11); RM' (15); MH55); M:U15). 

Remarks.— A\\ the Neotoma remains compare well with N. alhigula. The occlusal 
patterns for adult teeth are well-rounded as in A^. alhigula and A^. lepida (Desert wood 
rat), but the teeth are much larger than those of modem A^. lepida. The anterolingual 
re-entrant on the M, are very shallow as in A^. alhigula compared to the deep, microtine- 
like dental characters of A^. mexicana (Mexican wood rat). 

Five species of Neotoma presently live in Arizona (Colorado Plateau), more than 
in any other state. Only A^. lepida, N. alhigula, and A^. mexicana inhabit southern 
Arizona today, and only the last two presently occur in the vicinity of Deadman Cave. 
A^. alhigula lives in desert-grassland and desertscrub habitats while N. mexicana occurs 


Table 2. Measurements of modem and fossil dentaries (Deadman and Rampart caves) of Bassariscus 
astutiis. Measurements are rounded to the nearest 0.5 mm. 




Deadman Cave 

Modem Arizona 

Rampart Cave, Arizona (northem) 

Alveolar length P4-M, 







Deadman Cave 

Modem Arizona 

Rampart Cave, Arizona (northem) 

Alveolar length Mi 






in higher woodland and forest areas. Neotoma stephensi (Stephen's wood rat) occurs 
in the northem half of the state and A'', cinerea (Bushy-tailed wood rat) in the north- 
eastern sector (Cockrum 1982). The midden of the wood rat is found in numerous dry 
localities throughout the Southwest and is radiocarbon dated back to more than 40 000 
B.P. (Van Devender 1977, Van Devender and Spaulding 1979). 

Microtus species— Meadow vole 

Material.-L mandible (15043); LM, (2: 15044); RM, (15045); RM^ (3: 15046). 

Remarks.— We have not identified the fossil teeth to the specific level. We find it 
difficult to differentiate M. longicaudus (Long-tailed vole) from M. montanus (Montane 
vole) using isolated molars. Of the three complete fossil MjS examined, two had four 
closed alternating triangles and one had five triangles. 

None of the four species of vole found in Arizona presently occur near Deadman 
Cave. Microtus mexicanus (Mexican vole) presently occurs in the mountainous region 
of eastern Arizona but may have had a wider, more western distribution, based upon 
the present isolated occurrence of M m. hualpaiensis in northwestern Arizona, in the 
Late Pleistocene and/or Early Holocene (Hall 1981). The Mexican vole was also iden- 
tified in the fossil remains from Papago Springs Cave (Skinner 1 942). Microtus mon- 
tanus (Montane vole) has a predominantly northwestern distribution in the United 
States. Its nearest occurrence to Deadman Cave is in the Arizona Strip region of 
northernmost Arizona and in east-central Arizona. Microtus pennsylvanicus (Meadow 
vole) lives mainly in northem and eastem North America but approaches Arizona in 
northwestem New Mexico. The Meadow vole may have had a more southem, moun- 
tainous distribution in the Late Pleistocene or Early Holocene based upon an isolated 
modem population in northwestem Chihuahua, Mexico (Bradley and Cockrum 1968). 
Microtus longicaudus occurs through much of westem North America, including north- 
eastem Arizona. An isolated population presently lives in the Pinaleiio Mountains only 
60 km east of Deadman Cave. Based on present geographic distributions, this species 
seems most likely to have inhabited the mountains of the Basin-and-Range province 
of southeastern Arizona during the Late Pleistocene and Early Holocene. 

Order CARNIVORA-Camivores 

Family Procyonidae— Racoons, coatis, and ringtails 

Bassariscus a^rw/w^— Ringtail 

Material. -K mandible P.-., (15062); R maxilla (15315); LP^ (15316). 

Remarks.— The fossil Ringtail specimen compares well with modem specimens 
except that the M2 is not developed in the fossil specimen, but is replaced by a distinct 
depressional scar where the tooth was to have developed. The alveolar length from P4 
to M, (Table 2) is slightly longer in the Deadman Cave specimen than in modem 
specimens. The mandible from Deadman Cave does not seem to be similar to the rami 


Table 3. Measurements of dentaries (modem and Deadman Cave) of Spilogale putorius. Measurements 
are rounded to nearest 0.5 mm. 




Deadman Cave 

Modem Arizona 

Modem Nevada (northern) 

Alveolar length Pj-Mj 







Deadman Cave 

Modem Arizona 

Modem Nevada (northern) 

Alveolar length P^-M' 








described as B. sonoitensis from Papago Springs Cave (Skinner 1 942). Late Pleistocene 
and Holocene localities of the Ringtail are shown in Mead and Van Devender (1981). 
The Ringtail is widespread in rocky habitats in the desert grassland, and woodlands 
of the Southwest, and lives today in Deadman Cave. Numerous modem scats are 
located throughout the cave, especially near the entrance as demonstrated by the ac- 
cumulations of seeds, insects, and bones. 

Family Mustelidae— Weasels, skunks, and badgers 
Spilogale /7wror/"w5— Spotted skunk 

Ma/ma/. -L mandibles (3; 1 5047-1 5049); R mandibles (6; 1 5050-1 5055); L max- 
illa (15056); R maxillae (2; 15057-15058); L humerus (15059). 

Remarks.— T\\Q specimens from Deadman Cave are consistently larger than mod- 
em specimens of S. putorius (Table 3). The Spotted skunk is common throughout the 
Southwest and can be found in the Santa Catalina Mountains down to the lower desert 
mountain ranges. Elsewhere, fossil remains of the Spotted skunk have been recovered 
from Arizona (Skinner 1942), Califomia (Stock 1930), and New Mexico (Harris 1977). 

Mephitis macroura— Hooded skunk 

Material. -L mandibles (2; 15060); L maxilla (15061); L humerus (15317). 

Remarks.— The left mandible compares most favorably with that of Mephitis 
macroura, being smaller than in M. mephitis (Striped skunk) or Conepatus mesoleucus 
(Hog-nosed skunk) yet definitely larger than in Spilogale. All three skunks occur in 
southern Arizona, including the Santa Catalina Mountains. Mephitis macroura has its 
present northem distribution in southem Arizona and New Mexico (Hall 1981). We 
know of no other Late Pleistocene-Early Holocene record of this taxon. 

Family Felidae— Cats 
Felis coAzco/or— Mountain lion 

Material.-La (15063); LC, (15064); RC, (15065). 

Remarks.— AM the canines compared well with F. concolor rather than Panthera 
leo atrox (American lion) which has been recovered from Late Pleistocene age deposits 
of the nearby Murray Springs site (Haynes 1968, J. J. Saunders personal communi- 
cation). The canines were compared in size to those of modem Arizona Mountain lions 
and fossil specimens from Rancho La Brea, Califomia, El Durado, Colorado, Tule 
Springs, Nevada (Kurten 1973), and Rampart Cave, Arizona (Table 4). There were no 
discernible differences other than that the Late Pleistocene Felis concolor canines from 
Rancho La Brea may have been slightly larger than those from Deadman Cave. 

The Mountain lion occurs historically and paleontologically throughout the South- 
west, including the Santa Catalina Mountains (Hall 1981). Although it is usually found 
in woodlands and forest country, it also lives in mgged ranges well within the Sonoran 


Table 4. Length (L; anterior-posterior) and breadth (B; labial-lingual) range of measurements of canines 
in modem and Late Pleistocene-Early Holocene Mountain lion (Felis concolor). Recent specimens from 
Arizona (44, 654, 22785, and 23444; Department of Ecology and Evolutionary Biology, University of 
Arizona, Tucson). Measurements (in millimeters) of fossil specimens, other than from Deadman and 
Rampart caves, from Kurten (1976). 

n = 







Recent Arizona 
Deadman Cave, Arizona 
Rampart Cave, Arizona 
Rancho La Brea, California 
El Dorado, Colorado 
Tule Springs, Nevada 














Order PERISSODACTYLA- Odd-toed ungulates 

Family Equidae— Horses 

Equus species— Horse 

Material — 2nd phalanx (15027). 

Remarks. — This single element of a horse could not be identified to species. Re- 
mains of extinct species of Horse are common throughout the Southwest (Stock 1930, 
Mawby 1967, Lindsay and Tessman 1974, Harris 1977, Harris and Porter 1980, Kurten 
and Anderson 1980). Skinner (1942) identified E. conversidens and E. tau from Papago 
Springs Cave. 

Order ARTIODACTYLA— Even-toed ungulates 

Family Cervidae— Cervids 

Odocoileus species— Deer 

Material.-l.V2 {\5Q>66). 

Remarks.— The Deadman Cave specimen of Odocoileus could not be identified to 
species. Today in southern Arizona O. hemionus (Black-tailed deer) lives in the lowland 
habitats while O. virginiana (White-tailed deer) is found on the mountain tops. Fossil 
remains of both species of deer are widespread in North America (Kurten and Anderson 
1980), although only a few Late Pleistocene localities in Arizona contain remains of 
deer (Lindsay and Tessman 1974, Mead 1981). 

Family Bovidae— Bovids 

Euceratherium collinum-Shvub-ox 

Material. — L mandible (15318). 

Remarks.— The single specimen of the extinct Shrub-ox was identified by Walter 
Dalquest and Ernest Lundelius (Dalquest 1981, personal communication). The occlusal 
surface of the teeth were very worn, indicating an old individual. This specimen is the 
first record of the Shrub-ox in Arizona. Kurten and Anderson (1980) describe this 
bovid as a large, specialized grazer that probably lived in the lower foothills (like at 
Deadman Cave) rather than in the high, forested mountains. Euceratherium collinum 
is also known from Burnet Cave, New Mexico (Schultz and Howard 1935), but is not 
a common component of Late Pleistocene faunas in the Southwest (Kurten and An- 
derson 1980). 



Although there are numerous Late Pleistocene age localities in Arizona, especially 
in the southern portion (Lindsay and Tessman 1974), most of these sites are isolated 


finds in alluvial deposits containing Mammuthus jeffersoni (Jefferson's mammoth; =M. 
columbi, fide Kurten and Anderson 1980), Camelops species (camel), Equus species, 
or Bison species (bison). 

The most current work in Arizona concerning Late Pleistocene (Wisconsinan) age 
deposits comes from the well-preserved, radiocarbon dated wood rat middens, but 
these localities rarely contain the larger animals (Van Devender and Mead 1978, Mead 
198 1, Mead et al. 1983). The only cave faunas studied in southern Arizona are Papago 
Springs Cave (Skinner 1942) and Ventana Cave (Haury 1950). Ventana Cave (145 km 
west of Deadman Cave, Fig. 1) was excavated primarily for its abundant archaeological 
remains. Two lower units, Volcanic and Conglomerate, were deposited during the 
Wisconsinan glacial episode (Haury 1950). Papago Springs Cave (112 km south of 
Deadman Cave, Fig. 1) was excavated solely for its Late Pleistocene vertebrate fossils, 
which were laborously chiseled from brecciated layers along the walls and ceiling of 
the cave (Skinner 1942). 

The two cave faunas are very different in composition (Table 1). The fourteen taxa 
recovered from the lower units in Ventana Cave were all mammals, whereas a bird 
and 32 mammals were recovered from Papago Springs Cave. Only Canis latrans (=C. 
caneloensis\ Coyote), Lepus californicus, and Taxidea taxus (Badger) are shared in both 
faunas. Of the 64 taxa from Deadman Cave (5 amphibians, 25 reptiles, 12 birds, and 
22 mammals), only Nothrotheriops shastensis is shared with Ventana Cave and five 
species {Spermophilus variegatus, Thomomys cf. T. bottae, Neotoma albigiila, Antro- 
zous pallidus, and Spilogale putohus) are shared with Papago Springs Cave. Each cave 
is in a different physiographic setting in southern Arizona and had different modes of 
fossil accumulation. Deadman Cave is a limestone cave on a large mountain mass 
between two major river valleys surrounded by Sonoran and Chihuahuan desertscrub 
communities; its fauna was collected predominantly by small carnivores and raptors. 
Ventana Cave is a volcanic rockshelter in a small, low-elevation desert mountain range 
presently surrounded by the hot, dry Sonoran Desert. The open cave provided some 
shelter, easy access, as well as water from a spring. Papago Springs Cave was an open 
limestone cave in rolling oak woodland in the Canelo Hills of southeastern Arizona. 

The Late Wisconsinan age assignment of the deposition of the Volcanic and Con- 
glomerate units in Ventana Cave is presumably correct (Haury 1950, see Long and 
Muller 1981). The fossils from Papago Springs Cave are only broadly assigned to the 
Late Pleistocene, but for several reasons we suggest that the Papago Springs Cave deposit 
was formed prior to the last glacial maximum in the Wisconsinan (>22 000 B.P.). The 
chamber within the cave has changed configuration greatly since the initial deposition 
of the fauna. A great abundance of rock rubble (10 m thick) with bones has filled the 
cavern and become cemented. According to Skinner (1942) the cave was sealed off 
from the outside for a period of time, allowing settling and cementing of the fossils to 
take place. A third stage in the history of the cave reopened the cavern entrances 
permitting recent faunal accumulations and partial erosion of the fossil deposits. We 
feel that such an accumulation may have required a few ten's of thousands of years, 
placing the time of deposition sometime in the Middle or Early Wisconsinan glacial 


Five species of amphibians (all anurans) were identified from Deadman Cave 
(Table 1). Of these, only Bufo punctatus and Scaphiopus couchi were reported prior to 
this report from the Late Pleistocene and Early Holocene of Arizona. We report Bufo 
cf. B. woodhousei, Scaphiopus cf. 5. hammondi, and Rana sp. for the first time as 
fossils in Arizona, although they have been previously recorded from Late Pleistocene 
localities in California, Nevada, and New Mexico. Except for B. woodhousei, all anurans 
from the Deadman Cave fauna are found nearby the cave today. 

Reptiles are better known than the amphibians from Late Pleistocene-Early Ho- 
locene deposits in Arizona. We report 25 reptiles, including 13 species of lizards and 
1 2 species of snakes, from Deadman Cave. Of the lizards, Callisaurus draconoides, 


Holbrookia maculata, H. texana, Phrynosoma douglassi, P. modestum, P. solare, Sce- 
loporus cf. S. clarkii, and Urosaurus ornatus have not been reported previously from 
the Late Pleistocene of Arizona. The records of Heloderma suspectum from Vulture 
Cave, Arizona, and Gypsum Cave, Nevada, may be Early or Middle Holocene in age, 
and therefore this species perhaps should be added to the above list. Mdst of these 
lizards have previous fossil records from sites in California, Nevada, and/or New 
Mexico. Callisaurus draconoides is reported for the first time as a Late Wisconsinan- 
Early Holocene fossil. Phrynosoma modestum and C. draconoides do not presently 
inhabit the region of Deadman Cave nor the immediate valley. The closest occurrence 
of/*, modestum is in Chihuahuan desert-grassland 95 km east of the cave. Callisaurus 
draconoides occurs just west of Deadman Cave in lower Sonoran desertscrub. The other 
lizards from Deadman Cave, as well as most of the snakes, now live within a raptor's 
hunting range of the fossil locality. Of the 1 2 species of snakes identified from Deadman 
Cave, only Gyalopium canum has not been previously reported as a fossil from Arizona, 
and is out of its present distributional range at Deadman Cave. 

The birds from Deadman Cave represent only the fifth Late Pleistocene avifauna 
to be reported from Arizona. This contrasts markedly with the adjacent states of New 
Mexico and California, each of which has an excellent record of Late Pleistocene birds. 
The Deadman Cave avifauna is especially significant in that the diverse Sonoran Desert 
avifauna has yielded very few Pleistocene fossils. 

The Deadman Cave fauna contains several new avian fossil records, none of which 
is unexpected. Micrathene whitneyi and Catharus guttatus are reported for the first 
time as fossils, while Cyrtonyx montezumae is recorded paleontologically for the first 
time anywhere in the United States. New Arizonan fossil records include Colinus 
gambelli, Colinus species, Zenaida cf. Z. macroura, Otus species, Asio otus, the in- 
determinate caprimulgid, icterine, and Colaptes auratus. 

Mammals are the best known group of fossil vertebrates from Arizona. Twenty- 
two mammals are identified from Deadman Cave, most of which can be found in the 
Santa Catalina Mountains today or in the surrounding valleys. Exceptions are the three 
large extinct species {Equus species, Nothrotheriops shastensis, and Euceratherium col- 
linum) and Microtus species which lives 60 km to the east of Deadman Cave. 

Paleoenvironment of Deadman Cave 

The amphibian remains indicate that the region around Deadman Cave was at 
least as moist at the time of deposition as it is today. Certainly pools of water were 
nearby as they are today in portions of the Santa Catalina Mountains and along the 
major streams and rivers. The lizard fossils argue for either a community that was a 
composite of today's vegetational communities, or, a few vegetation groups abutting 
each other in close proximity to the cave. The presence of Phrynosoma modestum 
suggests that a desert-grassland area may have occurred on the broad flat hills above 
the large river valley, at the base of the mountain mass. Callisaurus draconoides, 
Holbrookia texana, Sceloporus magister, and Heloderma suspectum indicate a more- 
or-less open desertscrub to desert-grassland. Such areas could easily abut an open 
woodland. This community would probably be in the lower part of the bajadas between 
hills. The rest of the lizard fauna indicates several vegetation communities ranging 
from desert and desert-grassland to woodland. Certainly some areas of talus or rock 
outcrops were nearby. Most of the snake population from Deadman Cave suggests little 
to distinguish the region then from what it is today, although Gyalopium canum in- 
dicates a cool grassland habitat. 

Except for the possibly extinct icterid, the entire avifauna of Deadman Cave con- 
sists of living taxa that occur today in southern Arizona. No birds in the fauna are 
restricted to coniferous habitats. In fact, Colinus gambelli and Micranthene whitneyi 
argue strongly against any sort of adjacent coniferous forest. Colinus gambelli suggests 
the presence of desertscrub, while Cyrtonyx montezumae is characteristic of open oak 
woodlands, and pine-oak woodlands. Cyrtonyx is the only bird from Deadman Cave 


that is not found at least occasionally in desertscrub today, although Turdus migratorius 
and Catharus guttatus are certainly more abundant in wooded areas than in desertscrub, 
and never nest in the latter. The fossil birds from Deadman Cave suggest a slightly 
more grassy and wooded condition in the Latest Pleistocene and Earliest Holocene 
than today, such as an evergreen oak grassland mixed with desertscrub and desert- 
grassland on the more xeric exposures, and oak woodland on more protected, mesic 

The mammals generally indicate an open woodland with some areas more vege- 
tated and other areas more xeric and open. None of the mammals are restricted to 
forested habitats, although many of them do occur today in the forested higher ele- 
vations of nearby larger mountains. Notiosorex crawfordi, Microtus sp., Reithro- 
dontomys montanus, and Euceratherium collinum may argue for an open, possibly 
grassy woodland. Today this sort of habitat occurs just a few hundred meters upslope 
from Deadman Cave. Perognathus cf P. flavus, Dipodomys spectabilis, Thomomys cf 
T. bottae, and Sigmodon cf S. arizonae indicate that a desertscrub to desert-grassland 
was nearby. 

No distinctly boreal or mesic mammals (e.g., Sorex ornatus, Ornate shrew; S. 
palustrus, Water shrew; Ochotona princeps, Pika; Marmota flaviventris, Yellow-bellied 
marmot; Glaucomys sabrinus, Northern flying squirrel) were recovered from Deadman 
Cave. This may be because the faunal assemblage is of Late Wisconsinan-Early Ho- 
locene transition in age, and the boreal-mesic (full glacial) elements had already become 
extirpated locally. This may be the case with the Spermophilus lateralis from Ventana 
Cave. An alternative is that most of these elements never did occur in southern Arizona 
during the last Wisconsinan full and late glacial (assuming that the Papago Spring Cave 
deposit is Middle Wisconsinan or older in age). Presumably if any geographically or 
ecologically extralocal small animals, such as listed above, were contemporaneous with 
the Shasta ground sloth, Horse, and Shrub-ox, they too would have been observed in 
the Deadman Cave deposit. 

The Deadman Cave faunal record is very similar to those of Early Holocene-Late 
Pleistocene wood rat faunas in the Sonoran Desert (Van Devender and Mead 1978, 
Mead et al. 1983) and Grand Canyon (Van Devender et al. 1977, Mead and Phillips 
1981) in that the small vertebrates were conservative with few animals out of their 
present range, although there was a greater change in the local vegetation. The pollen 
record at Willcox Playa, 80 km east of the Santa Catalina Mountains, recorded a pine 
forest at 1 200 m elevation 20 000 years ago in an area that now supports desert-grassland 
(Martin 1963). This is an estimated lowering of the vegetation zones by about 1000 m 
elevation. The wood rat midden record for lower areas in the Sonoran Desert in 
southwestern Arizona recorded a complex elevational lowering of 600 m or less of 
certain woodland plants into the desert. An equivalent lowering of vegetation zones in 
the Santa Catalina Mountains would imply a shift from the modem desert-grassland- 
oak woodland vegetation to Pondersoa pine mixed conifer forest or a Mexican pine- 
oak woodland, depending on the distribution and abundance of precipitation in the 
Late Wisconsinan (Whittaker and Niering 1968). In the Early Holocene a Mexican 
pine-oak woodland was probably near the site until about 8000 years ago with the 
grassland developing more recently. Like the Microtus sp., a population of Abies 
lasiocarpa (Corkbark fir) presently on the top of the Santa Catalina Mountains, is also 
isolated from its nearest population in the spruce forests of the Pinalefio Mountains. 
Most of the remaining Deadman Cave fauna would be found in some sort of open 
woodland today. Several animals including Callisaurus draconoides, Heloderma sus- 
pectum, Dipodomys spectabilis, Phrynosoma modestum, and Colinus gambelli do not 
live in Mexican pine-oak woodland today and are more likely to occur in open desert- 
grassland or desertscrub and may represent the Early Holocene portion of the Deadman 
Cave deposit. Another possibility which has been demonstrated for a few animals in 
the Sonoran Desert (Van Devender and Mead 1978; Mead et al. 1983) is that under 
an equable Late Wisconsinan climate certain animals now confined to deserts were 
able to live in more diverse woodland communities. 



The vertebrate fauna of Deadman Cave includes 5 amphibians, 25 reptiles (13 
lizards and 12 snakes), 12 birds, and 22 mammals for a total of 64 species. The 
implication from this faunal assemblage is that by the end of the late glacial and the 
beginning of the post glacial (8000-12 000 B.P.), most of the local fauna was essentially 
as it is today— modem. Only one amphibian {Bufo woodhousei), three reptiles (Calli- 
saurus draconoides, Phrynosoma modestum, Gyalopium canum), and one mammal 
{Microtus species) are locally extirpated, although all still occur in southern Arizona. 
Animals unequivocally extinct are the Shrub-ox, Horse, and Shasta ground sloth, all 
large mammals. An unidentified icterine bird may prove to be an extinct species. 
Overall, the Deadman Cave fauna suggests that the vegetation community of the Late 
Pleistocene and Early Holocene were rather similar to those found today in the same 

The Deadman Cave bone assemblage has expanded our knowledge of the Late 
Pleistocene-Early Holocene fauna of southern Arizona, and has provided new questions 
on Late Pleistocene zoogeography of the hot deserts. Further stratified and datable cave 
deposits and wood rat middens need to be studied to refine when the faunas of southern 
Arizona became "modem." Upland localities need to be studied to determine whether 
boreal and/or mesic faunal elements ever existed in the Late Wisconsinan of southern 
Arizona, and whether the presently extralocal and local faunal assemblages ever co- 


Foremost we thank William Peachey, a caver from Tucson, Arizona, for finding 
the bone deposit in Deadman Cave and for realizing the cave's importance to vertebrate 
paleontology. Walter Dalquest and Ernest Lundelius provided the identification of the 
Shrub-ox, Greg McDonald verified the ground sloth identification; we appreciate their 
help. We thank Mary C. McKitrick for preliminary identification of certain avian fossils. 
Storrs L. Olson and the staff of the Division of Birds, United States National Museum 
of Natural History, Smithsonian Institution, made available their skeleton collection 
for comparative purposes. Our identification of the mammals was aided by the use of 
the mammalogy collection and the helpful suggestions provided by E. Lendell Cockrum 
and Yar Partzchian, Department of Ecology and Evolutionary Biology, University of 
Arizona, Tucson. The Grand Canyon National Park provided us with certain skeletons 
from their Study Collection for comparative purposes. Gene Hall helped prepare some 
of the Deadman Cave fossils. Austin Long of the Laboratory for Isotope Geochemistry, 
University of Arizona, Tucson, provided the radiocarbon date. Financial support was 
provided by grants from National Science Foundation to Paul S. Martin, University 
of Arizona (DEB75- 13944) and Thomas Van Devender (DEB76- 19784), and from the 
Smithsonian Institution (Predoctoral Fellowship, Scholarly Studies Program) to David 
Steadman. Emilee M. Mead drafted the figures and helped in various aspects of field- 
work. We thank Donald K. Grayson, two anonymous reviewers, and Gregory Pregill 
for critiquing and editing our manuscript. The Institute for Quaternary Studies provided 
final typing services. 

Literature Cited 

Armstrong, David M., and J. Knox Jones, Jr. 1 972. 

Notiosorex crawfordi. Mammalian Species No. 

Auffenberg, W. 1963. The fossil snakes of Florida. 

Tulane Studies in Zoology 10:131-216. 
Baker, Rollin H., and Karl A. Shump, Jr. 1978a. 

Sigmodonfulviventer. Mammalian Species No. 

, and . 1978/>. Sigmodon ochro- 

gnathus. Mammalian Species No. 97. 
Barbour, Roger W., and Wayne H. Davis. 1969. 

Bats of America. University Press of Kentucky, 

Baumel, J. J., A. S. King, A. M. Lucas, J. E. Breasile, 
and H. E. Evans. 1979. Nomina Anatomica 
Avium. Academic Press, London. 

Bradley, W. Glen, and E. Lendell Cockrum. 1 968. 
A new subspecies of the meadow vole (Micro- 
tus pennsylvanicus) from northewestem Chi- 
huahua, Mexico. American Museum Novi- 
tates No. 2325. 

Brattslrom, B. H. 1953a. The amphibians and 


reptiles from Rancho La Brea. Transactions of 
the San Diego Society of Natural History 1 1: 

— . 1953ft. Records of Pleistocene reptiles 
from California. Copeia 1953:174-179. 

— . 1954a. Amphibians and reptiles from 
Gypsum Cave, Nevada. Bulletin of the South- 
em California Academy of Science 53:8-12. 

— . 1954ft. The fossil pit-vipers (Reptilia: 
Crotalidae) of North America. Transactions of 
the San Diego Society of Natural History 1 2(3): 

— . 1964a. Amphibians and reptiles from cave 
deposits in south-central New Mexico. Bulletin 
of the Southern California Academy of Science 

— . 1 964ft. Evolution of the pit vipers. Trans- 
actions of the San Diego Society of Natural 
History 13(1 1):185-268. 
-. 1976. A Pleistocene herpetofauna from 

Smith Creek Cave, Nevada. Southern Califor- 
nia Academy of Science Bulletin 75:283-284. 

Brodkorb, Pierce. 1957. New passerine birds from 
the Pleistocene of Reddick, Florida. Journal of 
Paleontology 31:129-138. 

. 1964. Catalogue of fossil birds: Part 2 

(Anseriformes through Galliformes). Bulletin 
of the Florida State Museum, Biological Series 

1978. Catalogue of fossil birds. Part 5 

Bulletin of the Florida State Museum, Biolog- 
ical Series 23:139-228. 

Cockrum, E. Lendell. 1960. The recent mammals 
of Arizona. University of Arizona Press, Tuc- 
son, USA. 

. 1982. Mammals of the Southwest. Uni- 
versity of Arizona Press, Tucson, USA. 

Cole, Kenneth L., and Jim I. Mead. 1981. Late 
Quaternary animal remains from packrat mid- 
dens in the eastern Grand Canyon, Arizona. 
Journal of the Arizona-Nevada Academy of 
Science 16:24-25. 

Gehlbach, Frederick R., and J. Alan Holman. 1 974. 
Paleoecology of amphibians and reptiles from 
Pratt Cave, Guadalupe Mountains National 
Park, Texas. Southwestern Naturalist 19:191- 

Hall, E. Raymond. 1981. The mammals of North 
America. John Wiley and Sons, New York, 

Harris, Arthur H. 1977. Wisconsin age environ- 
ments in the northern Chihuahuan Desert: evi- 
dence from the higher vertebrates. Pp. 23-5 1 
in Roland H. Wauer and David H. Riskind 
(eds.). Transactions of the symposium on the 
biological resources of the Chihuahuan Desert 
region United States and Mexico. National Park 
Service Transactions and Proceedings Series 
No. 3. 

, and James S. Findley. 1964. Pleistocene- 
Recent fauna of the Isleta Caves, Bernalillo 
County, New Mexico. American Journal of 
Science 262:1 14-120. 

, and Linda S. W. Porter. 1980. Late Pleis- 

tocene horses of Dry Cave, Eddy County, New 
Mexico. Journal of Mammalogy 61:46-65. 
Haury, Emil W. 1950. The stratigraphy and ar- 

chaeology of Ventana Cave. The University of 
Arizona Press, Tucson, USA. 

Haynes, C. Vance. 1968. Preliminary report on 
the late Quaternary geology of the San Pedro 
Valley, Arizona. Arizona Geological Society, 
Southern Arizona Guidebook 111:79-96. 

Hill, William H. 1971. Pleistocene snakes from a 
cave in Kendall County. Texas. Texas Journal 
of Science 22:209-216. 

Holman, J. Alan. 1962. A Texas Pleistocene her- 
petofauna. Copeia 1962:255-261. 

. 1970. A Pleistocene herpetofauna from 

Eddy County. New Mexico. Texas Journal of 
Science 22:29-39. 

Hooper, Emmet T. 1952. A systematic review of 
the harvest mice (genus Reithrodontomys) of 
Latin America. Miscellaneous Publications of 
the Museum of Zoology, University of Mich- 
igan No. 77. 

Howard, Hildegarde. 1963. Fossil birds from the 
Anza-Borrego Desert. Los Angele County Mu- 
seum of Natural History, Contributions to Sci- 
ence No. 73. 

, and A. H. Miller. 1933. Bird remains from 

cave deposits in New Mexico. Condor 35:15- 

Hudson, Dennis M., and Bayard H. Brattstrom. 
1977. A small herpetofauna from the late 
Pleistocene of Newport Beach Mesa, Orange 
County, California. Bulletin of the Southern 
California Academy of Science 76:16-20. 

Jones, J. Knoxes, Jr., Dilford C. Carter, Hugh H. 
Genoways, Robert S. Hoffmann, and Dale W. 
Rice. 1982. Revised checklist of North 
American Mammals north of Mexico, 1982. 
Occasional Papers the Museum Texas Tech 
University 80:1-20. 

Kurten, Bjom. 1976. Fossil puma (Mammalia: 
Felidae) in North America. Netherlands Jour- 
nal of Zoology 26:502-534. 

, and Elaine Anderson. 1980. Pleistocene 

mammals of North America. Columbia Uni- 
versity Press, New York, New York. USA. 

Lindsay, Everett H., and Norman T. Tessman. 
1974. Cenozoic vertebrate localities and fau- 
nas in Arizona. Journal of the Arizona Acad- 
emy of Science 9:3-24. 

Long, Austin, and Paul S. Martin. 1974. Death 
of American ground sloths. Science 186:638- 

, and A. B. Muller. 1981. Arizona radio- 
carbon dates X. Radiocarbon 23:191-217. 

Lowe, Charles H. 1964. The vertebrates of Ari- 
zona. University of Arizona Press, Tucson, 

Martin, Paul S. 1963. Geochronology of pluvial 
Lake Cochise, southern Arizona, II pollen 
analysis of a 42-meter core. Ecology 44:436- 

. 1967. Prehistoric overkill. Pp. 75-120 m 

Paul S. Martin and Herbert E. Wright (eds.). 
Pleistocene extinctions, the search for a cause. 
Yale University Press, New Haven, Connect- 

Mawby, John E. 1967. Fossil vertebrates of the 
Tule Springs site, Nevada. Pp. 105-128 in H. 
M. Wormington and Dorothy Ellis (eds.). 
Pleistocene Studies in Southern Nevada. Ne- 


vada State Museum, Anthropological Papers 

Mead, Jim I. 1981. The last 30,000 years of faunal 
history within the Grand Canyon, Arizona. 
Quaternary Research 15:311-326. 

, and Arthur M. Phillips, III. 1981. The 

late Pleistocene and Holocene fauna and flora 
of Vulture Cave, Grand Canyon, Arizona. 
Southwestern Naturalist 26:257-288. 

, and Thomas R. Van Devender. 1981. Late 

Holocene diet of Bassariscus astutus in the 
Grand Canyon, Arizona. Journal of Mam- 
malogy 62:439-442. 

, Robert S. Thompson, and Thomas R. Van 

Devender. 1982. Late Wisconsinan and Ho- 
locene fauna from Smith Creek Canyon, Snake 
Range, Nevada. Transactions of the San Diego 
Society of Natural History 20:1-26. 
-, Thomas R. Van Devender, and Kenneth 

L.Cole. 1983. Late Quaternary small mam- 
mals from Sonora Desert packrat middens, Ar- 
izona and California. Journal of Mammalogy 

Meltzer, David J., and Jim L Mead. 1983. The 
timing of late Pleistocene mammalian extinc- 
tions in North America. Quaternary Research 

Miller, A. H. 1929. The passerine remains from 
Rancho La Brea in the paleontological collec- 
tions of the University of California. Univer- 
sity of California Publication, Bulletin of the 
Department of Geological Sciences 19:1-22. 

. 1932a. An extinct icterid from Shelter 

Cave, New Mexico. Auk 49:38-41. 

. 1 932i. The fossil passerine birds from the 

Pleistocene of Carpinteria, California. Univer- 
sity of California Publication, Bulletin of the 
Department of Geological Sciences 21:169-194. 

. 1947. A new genus of icterid from Rancho 

La Brea. Condor 49:22-24. 

. 1943. The Pleistocene birds of San Jo- 

secito Cavern, Mexico. University of Califor- 
nia Publication in Zoology 47:143-168. 

Montanucci, R. R., R. W. Axtell, and H. C. Des- 
sauer. 1975. Evolutionary divergence among 
collared lizards (Crotaphytus) with comments 
on the status of Gambelia. Herpetologica 3 1 : 

Mosimann, James E., and Paul S. Martin. 1975. 
Simulating overkill by Paleoindians. American 
Scientist 63:304-313. 

Phillips, A., J. Marshall, and G. Monson. 1964. 
The Birds of Arizona. University of Arizona 
Press, Tucson, USA. 

Rea, Amadeo M. 1980. Late Pleistocene and Ho- 
locene turkeys in the Southwest. Contributions 
to Science of the Natural History Museum of 
Los Angeles County 330:209-224. 

, and Lyndon L. Hargrave. In Press. Bird 

bones from Stanton's Cave, Arizona. 

Reeve, W. L. 1952. Taxonomy and distribution 
of the homed lizard genus Phrynosoma. Uni- 
versity of Kansas, Science Bulletin 34( 1 4):8 1 7- 

Schultz, B. C, and E. B. Howard. 1935. The fauna 
of Burnet Cave, Guadalupe Mountains, New 
Mexico. Proceedings of the Philadelphia Acad- 
emy of Natural Science 87:273-298. 

Skinner, Morris F. 1942. The fauna of Papago 
Springs Cave, Arizona and the Study of Stock- 
oceros: with three new antilocaprines from Ne- 
braska and Arizona. Bulletin of the American 
Museum of Natural History 80:143-220. 

Smith, N. M., and W. W. Tanner. 1^72. Two new 
subspecies of Crotaphytus (Sauria: Iguanidae). 
Great Basin Naturalist 32:25-34. 

Stebbins, R. C. 1966. A field guide to western 
reptiles and amphibians. Houghton Mifflin 
Company, Boston, USA. 

Stock, Chester. 1930. Rancho La Brea. Los An- 
geles County Museum of Natural History, Sci- 
ence Series No. 20. 

Thaeler, Charles S. 1980. Chromosome numbers 
and systematic relations in the genus Tho- 
momys (Rodentia: Geomyidae). Journal of 
Mammalogy 61:41 4-422. 

Thompson, Robert S., Thomas R. Van Devender, 
Paul S. Martin, Theresa Foppe, and Austin 
Long. 1980. Shasta ground sloth (A^c»//zro?/26'- 
riops shatense Hoffstetter) at Shelter Cave, New 
Mexico: environment, diet and extinction. 
Quaternary Research 14:360-376. 

Van Devender, Thomas R. 1973. Late Pleistocene 
plants and animals of the Sonoran Desert: a 
survey of ancient packrat middens in south- 
western Arizona. Ph.D. Dissertation. Univer- 
sity of Arizona, Tucson. 

. 1977. Holocene woodlands in the south- 
western deserts. Science 198:189-192. 

, and Richard D. Worthington. 1977. The 

herpetofauna of Howell's Ridge Cave and the 
paleoecology of the northwestern Chihuahuan 
Desert. Pp. 85-106 in Roland H. Wauer and 
David H. Riskind (eds.). Transactions of the 
symposium on the biological resources of the 
Chihuahuan Desert region United States and 
Mexico. National Park Service Transactions 
and Proceedings Series No. 3. 

, and Jim I. Mead. 1978. Early Holocene 

and late Pleistocene amphibians and reptiles 
in Sonoran Desert packrat middens. Copeia 

, and W. Geoffrey Spaulding. 1979. De- 
velopment of vegetation and climate in the 
Southwestern United States. Science 204:701- 

, Amadeo M. Rea, and Michael L. Smith. 

In Press. An interglacial vertebrate fauna from 
Rancho La Brisca, Sonora, Mexico. Transac- 
tions of the San Diego Society of Natural His- 
-, Arthur M. Phillips, III, and Jim I. Mead. 

1977. Late Pleistocene reptiles and small 
mammals from the Lower Grand Canyon of 
Arizona. Southwestern Naturalist 22:49-66. 

Wallace, Robert M. 1955. Structures at the north- 
em end of the Santa Catalina Mountains, Ar- 
izona. Unpublished Ph.D. Dissertation. Uni- 
versity of Arizona, Tucson. 

V/hittaker, R. H.,andW. A. Niering. 1965. Vege- 
tation of the Santa Catalina Mountains: a gra- 
dient analysis of the south slope. Ecology 46: 





%.: ^r 


Volume 20 Num6§i'^pp. 277-300 20 November 1984 

A Pliocene Flora from Chula Vista, 
San Diego County, California 

Daniel I. Axelrod 

Department of Botany, University of California, Davis, California 95616 USA 

Thomas A. Demere 

Department of Geology, San Diego Natural History Museum, P.O. Box. 1390, San Diego, California 921 12 

Abstract. A small fossil flora from the marine Upper Pliocene San Diego Formation suggests that 
the adjacent coastal plain was then covered with an avocado-Monterey pine-live oak woodland as- 
sociated with palm, cottonwood, willow, and sycamore along streams. Fossil digger pine apparently 
was confined to drier, warmer sites away from the coast. At higher, cooler levels farther inland were 
stands of fossil Jeffrey pine. Precipitation was near 50-58 cm over the lowlands, increasing to about 
65 cm in stands of fossil Jeffrey pine near 450-600 m. The fossil avocado, palm, and pine (afl!". Pinus 
radiata var. binata) indicate summer rainfall, consistent with the Late Pliocene higher-than-present sea 
surface temperature. Mean annual temperature on the coast was approximately 1 6°C, and annual range 
was about 7-8°C, equability was near M 70 with frost absent along the coast, light in the interior. 

Comparison with Pliocene floras in northern California shows that the Chula Vista flora lived in 
a separate floristic province, one corresponding with cismonlane southern California which has been 
a distinct floristic province since at least the Middle Miocene. Two new species of fossil pine are 
described: Pinus diegensis new species (allied with the living P. radiata var. binata) and P. jeffreyoides 
new species (similar to the living P. jeffreyi). 

The Chula Vista flora provides new evidence regarding the evolutionary history of Pinus radiata 
populations, and further insight into the disjunct distribution of taxa in the montane conifer forests of 
southern California and Baja California. 


The recent discovery of a small flora of Late Pliocene age (ca. 3 m.y.) at Chula 
Vista, in southwestern San Diego County, California (Fig. 1), provides preliminary 
information on the late Tertiary flora, vegetation, and climate of the region. The flora 
lived at a critical time in Neogene environmental history, one characterized by the 
cooler, moister climate that followed the warm, dry episode of the latest Miocene (5- 
6 m.y.) (Axelrod 1980Z?). The climatic transition, which heralds the build-up of ice 
sheets in Alaska and border areas, resulted in the displacement southward of relatively 
xeric sclerophyllous vegetation with numerous Madrean taxa. and their replacement 
by a more mesic flora. The nearest floras of comparable age are in central and northern 
California (Fig. 2). Analysis of the Chula Vista flora thus provides a general, though 
tentative, basis for interpreting the regional floristic differences that arose as cooler, 
moister climate spread southward and into lower altitudes in the Late Pliocene. 


The geology of the classical marine Pliocene San Diego Formation at San Diego 
has been summarized recently (Demere 1983). This rock unit crops out over a broad 
area which includes much of the southwestern portion of San Diego County, and the 
extreme northwestern comer of Baja California, Mexico. The San Diego Formation 


Late Cenozoic Floras 
of California 



t 1 1 T" 

50 100 150 200 Kilomelers 

Figure 1 . Late Cenozoic floras to which reference is made. 

was deposited during a marine transgression of the Neogene San Diego Basin, which 
hke other onshore sedimentary basins in southern California (e.g., Ventura Basin, Los 
Angeles Basin) is structurally related to the wrench and extensional tectonics of the 
continental borderland. Deposition began during the Late Pliocene and apparently 
continued into Early Pleistocene time, accumulating at least 75 m of marine and 9 m 
of nonmarine sedimentary rocks. The overall stratigraphic sequence suggests a succes- 
sive shallowing and filling of this basin. It is now apparent that extensional tectonics 
have controlled both the initial deposition as well as the present outcrop distribution 
of this rock unit. Numerous high-angle normal faults striking north to northwest cut 
the area into a series of small fault blocks which expose different portions of the Pliocene 

In an attempt to correlate between the fault blocks, Demere (1983) informally 
subdivided the San Diego Formation into a "lower" and an "upper" member using 















Seac 1 i f f 




S oboba 


, g 



^ -H 




>~ 3- 

. 2 

Chula Visf a - 

Santa Clara 




Santa Rosa 



C al i n g a 



Turlock Lake 

Broken Hill 


^ 0- 





Mount Eden 


Lower Petaluma 





Mulholland — 





Pi r u Gorge 

Jacal i t OS 







/ — 


Figure 2. Ages of Late Cenozoic floras in California. 

both lithologic and paleontologic criteria. The "lower" member is characterized by 
yellowish, very fine-grained, micaceous, massive, friable sandstones with locally well- 
bedded sequences of laminated and cross-bedded sandstones, pebble-to-cobble con- 
glomerates and well-cemented shell beds. This "lower" member is richly fossiliferous 
and has produced the bulk of the marine invertebrate fauna so well known through 
the work of Grant and Gale (1931) and Hertlein and Grant (1944, 1960, 1972). In 
addition, the diverse avifauna (Howard 1949, Miller 1956) and cetacean assemblage 
(Barnes 1973, 1 976) reported from the San Diego Formation have been largely collected 
from the "lower" member. 

Lithologically the "upper" member is characterized by well-bedded sequences of 
pebble-to-cobble conglomerate containing reworked "Poway" clasts, well-cemented 
fossiliferous sandstones, and medium- to coarse-grained friable sandstones. Marine 
invertebrate fossils are locally common in this member. 

In terms of paleoenvironment, the "lower" member contains a middle-to-outer- 
shelf molluscan fauna characterized by Patinopecten healeyi, Pecten stearnsii, Luci- 
noma annulata and Opalia varicostata. In contrast, molluscs from the "upper" member 
indicate deposition in littoral-to-inner-shelf depths. Characteristic species include Pec- 


ten bellus, Argopecten hakei, Niicella lamellosa and Dendraster ashleyi. Both members 
reflect normal marine deposition in a broad coastal embayment probably similar to 
present-day Monterey Bay along the central California coast. 

Barnes (1976) correlated the San Diego Formation ("lower" member) with the 
Blancan North American land mammal stage. This was based on the occurrence of the 
horse Equus, and is supported by the recent discovery at Chula Vista of teeth referable 
to the bunodont mastodon {Stegomastodon rexroadensis Woodbume). Relying on the 
stratigraphic ranges of moUuscan species the "lower" member is correlative with the 
upper portion of the San Joaquin Formation in the San Joaquin Basin, the Careaga 
Formation in the Santa Maria Basin, and the Niguel Formation and the upper Fernando 
Formation in the southeastern Los Angeles Basin, all Late Pliocene in age (Woodring 
and Bramlette 1951, Vedder 1972). In turn the "upper" member of the San Diego 
Formation is correlated with the lower Santa Barbara Formation in the Ventura Basin 
which is considered to be Late Pliocene to Early Pleistocene in age (Keen and Bentson 
1944, Lajoie et al. 1982). The meager microfossil record available in the San Diego 
Formation (Ingle 1967, Mandel 1973) suggests that it is apparently no older than 
planktonic foraminiferal zone N-21 (approximately 3.0 million years B.P. Late Plio- 
cene) and is perhaps as young as the Emiliania annula calcareous nannoplankton 
subzone (approximately 1.5 million years B.P., Early Pleistocene). 

The section at Chula Vista consists of approximately 73 m of fossiliferous San 
Diego Formation. Here the Pliocene marine rocks rest disconformably on fluvial and 
lacustrine sedimentary rocks of the Middle Miocene Sweetwater Formation. Overlying 
the San Diego Formation along an irregular erosion surface are fluvial and alluvial 
sedimentary rocks of the Lower Pleistocene Sweitzer Formation (=Lindavista For- 
mation of Kennedy 1975). The lower 46 m of the Chula Vista Pliocene section contain 
marine invertebrate taxa characteristic of the "lower" member of the San Diego For- 
mation. This part of the section is also characterized by locally common marine ver- 
tebrate fossils and has produced all of the paleobotanical material described in this 

In terms of general stratigraphic position, the described fossil pine cones were 
found betwen 24 and 30 m above the base of the San Diego Formation, the fossil leaves 
between 12 and 19 m, and a few poorly preserved shipworm {Teredo)-bovQ6. logs 
between 12 and 21 m. 

The fragmentary cones occur in large, elongate (up to 40 cm in diameter), limy 
concretions which formed around large bones of fossil baleen whales. Steinkems of 
marine invertebrates also occur in these concretions. The cones are preserved as natural 
internal and external molds and vary from incomplete three-dimensional specimens 
to only partial imprints. 

Fossil leaves occur as dense concentrations of flat-lying and stacked leaf material 
in both fine-grained sandstone and sandy siltstone. The leaves are preserved as iron- 
stained imprints and are largely fragmentary. The leaves were collected from thin ( 1 5 
cm thick) sandstone and siltstone strata interbedded with fossiliferous sandstones con- 
taining typical "lower" San Diego Formation marine invertebrate taxa (e.g., Patino- 
pecten healeyi and Lucinoma annulatd). The occurrence of terrestrial plant material in 
this marine setting points to offshore transport, no doubt first under fluvial and then 
under marine conditions. The concentration of leaf material in thin stratigraphic ho- 
rizons suggests marine transport by storm-generated debris flows. Otherwise, it seems 
doubtful that they would be concentrated in a local area. The cones probably were 
transported in a similar manner. The cone fragments are the result of partial preser- 
vation in the concretions, the remaining portion of the cone was not preserved in the 
soft compacted sandstones that enclose the concretions. 

Not described in this report, but occurring within the Pliocene section at Chula 
Vista, were a number of severely shipworm-bored, calcareously cemented logs. Some 
of these logs were up to 3 m in length and had broken branch stems. They all were 
lying parallel to the bedding upon locally pebbly fossiliferous sandstone. 


Table 1. Systematic list of floral species from the San Diego Formation, Chula Vista, California. 

Pinaceae Salicaceae 
Pinus diegemis n. sp. Populus alexanderi Dorf 

Finns jejfreyoides n. sp. Salix wildcatensis Axelrod 

Pinus pieperiDod T^^g^c^^^ 

Arecaceae Quercus lakevillensis Dorf 

Gen. et sp. indet. (Palm) Lauraceae 

Juncaceae Persea coalingensis (Dorf) Axelrod 

Gen. et sp. indet. (Reed) Platanaceae 

Platanus paucidentata Dorf 


The small paleobotanical collection from Chula Vista includes ten species distrib- 
uted among three conifers, two monocotyledons, and five dicotyledons. Two of the 
pines are new. The monocotyledons are represented by specimens too incomplete to 
refer to species. 

The fossil taxa are represented by only a few specimens each, and most are frag- 
mentary, reflecting their transport into the offshore. Fifteen specimens are referred to 
Persea. Of the remainder, Pinus diegensis is represented by 5 specimens, Populus 4, 
Platanus, Salix, Pinus Jeffrey oides 3 each, and Pinus pieperi, palm, Quercus and reed 
by one each. 

All of the fossils are similar to living taxa and have been assigned to fossil species 
following nomenclatural convention (Axelrod, 1980Z?:205). As judged from their re- 
lationships, the flora is composed of 8 trees, one shrub and one herbaceous perennial. 

An understanding of the ecology of extant species closely allied to the fossil plants 
in the Chula Vista flora provides a basis for reconstructing the paleovegetation and 
climate of the region. These allied modem species can be separated into four ecologic 
and geographic groups. 

First, several of the allied living species are widely distributed in central and 
southern California. These include sycamore {Platanus), cottonwood {Populus) and 
willow {Salix). Today, these trees inhabit streambanks in the coastal area from near 
San Francisco Bay southward, reaching also into the interior. The oak {Quercus agri- 
folia) forms dense woodlands in moister, more equable sites near the coast and con- 
tributes to open oak-grass communities at low to moderate elevations in more inland 
areas of the outer Coast Ranges. Quercus is also found frequently along stream margins 
where it is associated with cottonwood, sycamore, and willow. The fossil cottonwood 
{Populus alexanderi) is similar to P. trichocarpa var. trichocarpa that has roundish- 
ovate leaves and lives near the coast in mild equable climate from the San Francisco 
Bay region southward. It differs from the black cottonwood that occurs in the mountains 
of California and northward, which has larger, ellipitic-ovate leaves and is best referred 
to P. hastata Dode. 

Second, digger pine {Pinus sabiniana) characterizes the pine-oak woodland-grass 
vegetation of the inner Coast Ranges and lower slopes of the Sierra Nevada. It reaches 
the margin of southern California in Santa Ynez Valley north of Santa Barbara and on 
Liebre Mountain several miles southeast of Tejon Pass. It only approaches the 
coast in the Santa Lucia Mountains near Gorda where it seems to be a relict of the 
Xerothermic period. This also appears to account for its disjunct occurrence in Santa 
Ynez Valley. In the south Coast Ranges it inhabits flats adjacent to rivers, notably the 
Salinas, Santa Maria, Santa Ynez, and Sisquoc, which at times of flood probably 
transport its large cones to the coast. Among its associates in riparian sites are species 
of cottonwood, willow, sycamore, and live oak similar to taxa in the Chula Vista flora. 

Third, Pinus jeffreyi, allied to the fossil P. jeffreyoides new species, now occurs in 


Figure 3. Guadalupe Island pine {Pinus radiata var. binata) and palm {Brahea edulis) on Guadalupe Island 
have allied laxa in the Chula Vista fossil flora. (Photo by Reid Moran.) 

the mountains of eastern San Diego County. The stand nearest to Chula Vista is 55 
km northeast at Pine Valley, at an elevation of 1050 m. There it is associated with live 
oak (Quercus agrifolia). Streams in the area also support species of willow, cottonwood 
and sycamore similar to those in the fossil flora. The bordering slopes are covered with 
dense chaparral composed of Adenostoma, Arctostaphylos, Ceanothus, Cercocarpus, 
Quercus, and others. It is to be noted that P. jejfreyi replaces P. ponderosa as the lowest 
montane conifer forest species in the southern Peninsular Ranges. Here it is a distinct 
ecotype as compared with P. jejfreyi in the Sierra Nevada which is a member of the 
red fir-white pine-mountain hemlock forest that is snowbound all winter. Only an 
occasional light snow occurs at its lower elevations in San Diego County and in the 
Sierra Juarez and Sierra San Pedro Martir, northern Baja California, where it also meets 
oak woodland and chaparral vegetation, generally at levels near 1400-1500 m (see 
Nelson 1922). 

Fourth, three Chula Vista fossil species are allied to living taxa in western Mexico. 
Avocado (Persea), which was abundant in California into the close of the Tertiary 
(Sonoma, Wildcat, Turlock Lake, Broken Hill, Coalinga floras), now occurs from south- 
em Sonora southward in the Sierra Madre Occidental, and other species are in the 
eastern United States, chiefly along the Atlantic seaboard. In southern Sonora and 
Sinaloa, Persea is a frequent member of pine-oak woodland vegetation, associated with 
sycamore, willow, palm and other riparian taxa in valley bottoms. It occupies sheltered 
sites at 1000-1200 m under mild, equable climate (Gentry 1942, 1946). 

Cones of Pinus diegensis new species are similar to those of P. radiata var. binata 
of Guadalupe Island, situated 350 km off' the central coast of Baja California at 29°N 
latitude. This pine is confined chiefly to the middle-upper slopes of the island, generally 
at altitudes above 800 m where it is in a persistent fog belt during summer. The 
occurrence of P. diegensis on the mainland as recently as 3 m.y. ago suggests that the 
living P. radiata var. binata did not originate in insular isolation but probably was 
confined to Guadalupe Island as drier climate spread over the region. 

A fragmentary specimen of palm frond from Chula Vista shows relationship to 



Figure 4. Present environment in the area of the Chula Vista fossil flora. 

Sabal and also to Brahea (^Erythea). Sabal is a frequent member of the pine-oak 
woodland in the Sierra Madre Occidental of Sonora where it occurs along streams with 
avocado, willow, sycamore and other riparian inhabitants. Of considerable interest is 
the presence of Brahea edulis H. Wendl. ex. S. Wats, on Guadalupe Island where it 
occurs in deep, moist canyons with Pinus radiata var. binata (Howell 1941Z); Libby et 
al. 1968), as seen in Fig. 3. 


The community relations of modem species most similar to the fossil plants suggest 
that during the Late Pliocene this portion of the coastal strip was covered with dense 
woodland vegetation in contrast to the open coastal sage of today (Fig. 4). The abun- 
dance of avocado leaves in the collection implies that Persea dominated stream valleys 
and sheltered slopes near the shore. On moister slopes it was associated with fossil 
Guadalupe pine and probably formed a dense forest. Live oak was regularly associated 
with the pine and avocado. Along stream borders these trees occurred with sycamore, 
Cottonwood, willow, and palm. Toward the interior, fossil Guadalupe pine {P. diegensis) 
and live oak probably contributed to a more open woodland-grass community that 
gradually merged with a fossil digger pine association that included avocado and palm 
in sheltered sites, as well as riparian taxa. Fossil digger pine probably attained optimum 
development on exposed, hotter and drier south-facing slopes provided by granitic and 
metamorphic basement rocks farther inland. At higher, cooler and moister levels farther 
east were stands of fossil Jeffrey pine that reached down to a lower altitude than at 
present because precipitation was higher. 


Judging from conditions under which allied living taxa occur, annual precipitation 
at sea level during the Late Pliocene probably was near 58-63 cm as compared with 



^ ' -<^ I I I I I I I I I r 

^-! I I L 


Figure 5. Estimated paleotemperature for the Chula Vista coastal strip (A), the area of fossil digger pine 
(B), and the fossil Jeffrey pine forest (C). The difference in mean annual temperature between Chula Vista 
today and the inferred paleotemperature (circled area. A) is approximately 0.5°C. The radiating lines of 
warmth (W) indicate growing seasons defined by the number of days (d) with mean temperature warmer 
than the specified temperature (i.e., W 14°C has 183 days with mean temperature warmer than 14°C). The 
arcs M 70 and M 60 represent a measure of equability which decreases in all directions from T 14°C and A 
0°C. For further data see Bailey (1960, 1964). 

22 cm at Chula Vista today. Precipitation increased to about 70 cm in hills to the east 
at the lower margin of fossil Jeffrey pine forest. The remaining taxa in the fossil flora 
could readily survive under these rainfall totals. Fossil avocado, palm and Guadalupe 
pine (var. binatd) indicate summer rainfall. This resulted from a warmer sea-surface 
temperature than that of today, as shown by the marine macro-invertebrate fauna of 
the San Diego Formation. Hertlein and Grant (1954) estimated on the basis of the 
molluscan taxa that winter sea surface temperature during the Late Pliocene was similar 
to that now near Cedros Island (ca. 17-18°C), or fully 3-4°C higher than that in the 
San Diego area today (U.S. Navy 1956-58; U.S. Weather Bureau 1938). With warmer 
water, hurricanes from the eastern Pacific may have moved farther up the coast than 
they now do. In this regard, the summer of 1 983 in southern California was characterized 
by warmer-than-usual water offshore and increased warm season rainfall onshore. 

Because of the mixture of taxa from both coastal and inland sites, a single estimate 
cannot represent the thermal conditions for the entire fossil flora. As for the coastal 
strip, Guadalupe pine occurs at altitudes generally above 800 m. To judge from its 
occurrence in a summer fog belt, mean annual temperature is approximately 16°C as 
inferred from sea surface temperature (U.S.D.A., Climatic data); the persistent fog belt 
implies a low range of temperature, probably not more than 8-1 0°C. Because the marine 
invertebrate fauna of the "lower" San Diego Formation indicates that sea surface 
temperature was warmer than at present, temperature along the coastal plain was also 
warmer than at present, probably with a mean temperature near 1 6°C. This is suggested 
also by the abundance of avocado (Persea) specimens. Since avocado does not now 


produce fruit well in areas much north of Santa Barbara (Fig. 5), temperatures near 
there probably represent an extreme minimum for the fossil flora. Digger pine occurs 
today in Santa Ynez Valley (Cachuma Lake), not far from avocado groves, where mean 
temperature is about 16°C and the range is 10-1 1°C. Farther north in the Coast Ranges, 
as at Santa Margarita (alt. 300 m), mean annual temperature is 14.5°C and the annual 
range is 16°C. The palm, Guadalupe pine, and avocado suggest that fossil digger pine 
lived under conditions milder than those now at Santa Margarita, and probably were 
like those in the Cachuma Lake area. As for the interior, Jeffrey pine now occurs in 
Pine Valley where mean annual temperature, as estimated from that at Cuyamaca, 16 
km north and 350 m higher, is near 12.5°C and the mean monthly range is about 17°C. 
These estimates suggest that during the Late Pliocene the temperature along the coast 
was about 0.5-1.0°C warmer than that presently at Chula Vista. 

In view of the greater range of mean monthly temperature in the interior where 
fossil Jeffrey pine lived, mean annual temperature there probably was about 2-2. 5°C 
lower than on the coast (Fig. 5). This suggests a depression of regional climate of about 
300-400 m, and hence a lower altitude for taxa representing upland vegetation as 
compared with their present occurrence. Coupled with higher rainfall, under the pos- 
tulated paleotemperature fossil Jeffrey pine may have reached down to near 600 m as 
compared with 1050 m in Pine Valley today. Fossil Jeffrey pine would therefore have 
lived closer to the marine embayment, in sites from which its cones would more likely 
be transported seaward, especially during ffoods resulting from hurricanes that pre- 
sumably were of more frequent occurrence. As suggested in Fig. 5, equability on the 
coast was near M 70 as compared with M 67 at Chula Vista today. Frost was absent 
on the coast where the growing season probably had a mean daily temperature warmer 
than 15.2°C on 218 days of the year. Moderate light frosts might well be expected over 
the interior where fossil Jeffrey pine lived. The estimated paleotemperatures are shown 
by the circled areas A, B, and C, respectively, for the immediate coast, the central 
digger pine woodland, and the lower margin of Jeffrey pine forest (Fig. 5). 

Regional Comparisons 
Pliocene Floras 

The coastal oak-avocado-palm-pine woodland, the interior digger pine-oak wood- 
land and the upland Jeffrey pine forest of the Pliocene Chula Vista ffora differ consid- 
erably from contemporaneous vegetation in central and northern California (Fig. 1). 

The ffora from the Sonoma Formation at Neer's Hill, Santa Rosa (Axelrod 1944c) 
represents a coast conifer forest with redwood (Sequoia), lowland fir (Abies cf grandis), 
Douglas fir (Pseudotsuga cf menziesii), coast hemlock (Tsuga cf heterophylla) and 
winged seeds of weeping spruce (Picea cf brewehand). Associated with the conifers, 
and also forming a dense, broadleaved sclerophyll forest on warmer slopes, were species 
of oak (Quercus cf chrysolepis), chinquapin (Castanopsis [Chrysolepis] chrysophylla), 
California laurel (Umbellularid), and tanoak (Lithocarpus). Riparian species included 
sycamore, cottonwood, avocado, and willow. A few Tertiary relicts are recorded, no- 
tably species of Ilex, Persea, Trapa and Ulmus. It is estimated that the area received 
fully 89-100 cm of rainfall, some in summer, and that temperatures were cool though 
frost was absent. 

Representatives of the Sonoma ffora apparently extended farther north, as suggested 
by a small flora from the upper part of the Wildcat Group near Garberville (Axelrod 
1944c: 187, Dorf 1930). This fossil flora is composed chiefly of riparian taxa and is 
characterized by an absence of typical forest species. The flora apparently accumulated 
on a broad floodplain some distance from forests that occupied slopes bordering the 
lowland marine basin. The only forest representatives are logs of Sequoia and Pseu- 
dotsuga and a leaf of Ulmus. Members of floodplain vegetation included alders (Alnus 
cf. rhombifolia, rubra), avocado (Persea cf borbonia), sycamore (Platanus cf racemosa), 
black cottonwood (Populus cf hastata), willow (Salix cf lasiolepis) and California 
laurel ( Umbellularia). Some of these appear to represent Miocene relicts, notably the 


Table 2. Comparison of Chula Vista flora with related living taxa. 

Fossil species 

Allied living species 

Pinus diegensis 
Pinus jeffreyoides 
Pinus pieperi 
Platanus paucidentata 
Populus alexanderi 
Persea coalingensis 

Quercus lakevillemis 
Arecaceae sp. 


Salix wildcatensis 
Herbaceous perennial 

Juncaceae sp. 

P. radiata var. binata Englemann 

P. Jeffrey i Murray ^ 

P. sabiniana Douglas 

P. racemosa Nuttall 

P. trichocarpa Torrey & Gray 

P. podadenia Blake; P. borbonia 

(Linne) Sprengel 
Q. agrifolia Nee 
Braheal, Sabal? 

S. lasiolepis Bentham 


Persea, Populus, Ulmus and possibly the Platanus which is represented by an incomplete 
specimen. Their persistence here in the late Tertiary is understandable in view of their 
coastal position where there was high rainfall and low evaporation. It is evident that 
the Garberville flora shows little relation to the Chula Vista flora which is expectable 
in view of its position fully 1025 km southeast. 

The Napa flora (Axelrod 1950a), situated 45 km southeast of the Sonoma flora at 
Santa Rosa, represents a pine (Pinus cf. ponderosa)-Doug\as fir (Pseudotsuga) forest 
living near sea level. Forest associates included chinquapin (Castanopsis [Chrysolepis] 
cf. chrysophylla, C. sempervirens), Oregon grape (Mahonia cf. nervosa), ocean spray 
(Holodiscus cf. discolor), goldcup oak (Quercus chrysolepis), interior liveoak (Q. cf 
wislizenii), and California laurel (Umbellularia cf. californica). Sequoia is rare, with 
only three small twigs represented in the sample of over 700 specimens. Warmer slopes 
were covered with broadleaved sclerophyll vegetation composed of Castanopsis, Quer- 
cus, Umbellularia, and probably Persea. An open oak woodland-grass of coast liveoak 
(Quercus cf. agrifolia), valley oak (Q. lobata), and interior liveoak (Q. wislizenii) oc- 
cupied drier slopes. The warmer, driest sites supported a chaparral of whitehom 
(Ceanothus cf. leucodermis), mountain mahogany (Cerocarpus cf betuloides, C. cf. 
ledifolius), and toyon (Heteromeles cf. arbutifolia), though these shrubs also occurred 
in the oak woodland. Stream- and lake-border sites supported avocado (Persea), syc- 
amore (Platanus), willows (Salix cf. laevigata, S. scouleriana) and California laurel 
( Umbellularia). The assemblage reflects a drier, more continental climate than that in 
the Santa Rosa area to the northwest. The only relationship with the Chula Vista flora 
is seen in a few riparian taxa that ranged widely and through several vegetation zones. 

The Turlock Lake flora, slightly older than the Chula Vista, accumulated on the 
lowest floodplain in front of the central Sierra Nevada. It is preserved in the upper part 
of the Mehrten Formation 35 km east of Modesto (Axelrod 1980^?). The lake border 
assemblage included avocado (Persea), sycamore (Platanus), paloblanco (Forestiera), 
willow (Salix), and California laurel ( Umbellularia). The bordering slopes were covered 
with oak woodland-grass (Quercus cf. douglasii, Q. wislizenii) and scattered shrubs, 
including scrub oaks (Quercus cf. dumosa, Q. dunnii), ceanothus (Ceanothus cf sore- 
diatus), cofleeberries (Rhamnus californica, R. ilicifolia), and poison oak (Toxicoden- 
dron) that may have formed local brushy patches on exposed drier slopes. Cooler, 
moister sites supported broadleaved sclerophyll taxa, notably madrone (Arbutus), in- 
terior live oak (Q. wislizenii), and California laurel (Umbellularia). Reaching down 
from higher levels along stream valleys were members of a moister flora, including 
pine (Pinus cf. ponderosa) and smilax (Smilax cf. californica). Two exotics are in the 
flora, an aspen (Populus cf. tremula) and a cherry (Prunus), both of Asian affinity. 
There was some summer rainfall and annual precipitation totalled approximately 63 


cm as compared with 36 cm today. The flora shows Httle relationship with that at 
Chula Vista apart from several widely distributed riparian species. 

The small Coalinga flora from the upper San Joaquin Formation 10 km south of 
Coalinga indicates that the same general type of floodplain vegetation preserved at 
Turlock Lake ranged across the Central Valley. The fossils occur in sandstones of 
fluviatile origin interbedded with marine strata containing molluscs that represent the 
Pecten coalingensis zone (Dorf 1 930). The flora has abundant leaves of avocado (Persea) 
and sycamore {Plat anus), and together with cotton wood (Populus) and hackberry (Cel- 
tis) are indicative of riverbanks and moist sites. Adjacent interfluves were covered with 
oak (Quercus cf. douglasii) as well as silk-tassel bush (Garrya cf. elliptica). Further 
collecting at this site should provide a better representation of the flora and vegetation 
of this area. 

An indication of the flora that occupied the region slightly earlier (ca. 5 m.y.) is 
provided by a flora from Broken Hill, 30 km southeast, at the south end of North 
Dome, Kettleman Hills, in the basal part of the San Joaquin Formation (Axelrod 1 980^). 
It also provides evidence of vegetation in the area prior to uplift of the Coast Ranges 
directly west, an event that brought a drier, semidesert climate to this area. The flora 
occurs in sandstones of fluviatile origin that grade upward into marine beds. Floodplain 
vegetation included alder (Alnus), sycamore (Platanus), cottonwood (Populus), poplar 
(Populus euphmtica), aspen (Populus cf. tremula), California laurel (Umbel- 
lularia) as well as several willows (Salix cf. exigua, S. lasiandra, S. laevigata, S. lasi- 
olepis), and soapberry (Sapindus). Avocado (Persea) is abundant, magnolia is present, 
and both contributed to the floodplain vegetation. Well-drained interfluves were cov- 
ered with oak woodland-grass that included three oaks (Quercus cf. agrifolia, Q. doug- 
lasii, Q. wislizenii). There is also evidence of broadleaved sclerophyll vegetation, prob- 
ably on higher slopes to the west. These included tanoak (Lithocarpus), morheus oak 
(Q. morheus), and California laurel (Umbellularia), as well as avocado and magnolia. 
The flora has several exotics in addition to Magnolia and Persea, notably Populus cf. 
tremula, P. cf. euphratica, Sapindus and Ulmus, all indicating summer rainfall. Apart 
from these, the flora resembles vegetation of the lower middle slopes of the Santa Lucia 
Mountains 120 km west. Rainfall is estimated to have been near 75 cm annually, and 
climate was more equable than that now in this semi-desert region where precipitation 
totals 12 cm yearly. 

Evidently the Pliocene floras of central and northern California represent vegetation 
quite different than that at Chula Vista. Much as the vegetation of these areas differs 
today, the differences reflect the uniqueness of the floristic provinces represented. The 
principal links with the floras to the north were riparian taxa that ranged widely into 
very different vegetation zones and which have a broad time span. The transition from 
the northern coast conifer forests to the coastal vegetation of the Chula Vista area 
probably corresponded generally with the axis of the present Transverse Ranges. 

Plio- Pleistocene 

The preceding late Neogene floras were succeeded by those that represent colder, 
moister climates. This is illustrated by the Soboba flora from interior southern Cali- 
fornia (Axelrod 1967a), dated by the Bautista mammalian fauna at 1 m.y. This flora 
shows that mixed conifer forest reached down fully 1000 m below its present level in 
the San Jacinto Mountains which now tower above the fossil locality. Included as 
macrofossils that represent montane conifer forest are Abies concolor, Calocedrus de- 
currens, Pinus ponderosa, P. lambertiana, and Populus tremuloides. Associates included 
species of upper woodland and chaparral vegetation. Precipitation was near 70 cm 
annually as compared with 33 cm in the area today and mean annual temperature was 
fully 5.5°C lower than at present (Axelrod 1976: fig. 5). A flora of comparable age is 
represented in the Santa Cruz Mountains near Saratoga (Dorf 1930). It records forest 
taxa (Libocedrus, Pinus cf. lambertiana, Pseudotsuga) at much lower altitudes than 
occur near sea level in this area today. By contrast, the fossil site supports oak woodland- 


grass and broadleaved sclerophyll forest. Redwood forest is now at higher levels, and 
is not represented in the fossil flora. 

The magnitude of the climatic shift along the coastal strip in the Early Pleistocene 
is indicated by a flora from the marine upper Pico Formation (Early Pleistocene, 1 
m. y.), situated near Seacliff'on the coast west of Ventura (Axelrod 1983). Several cones 
of Douglas fir {Pseudotsuga menziesii) are represented, a species that now has scattered 
populations in the Santa Lucia Mountains 325 km north but has its principal southern 
area in the Santa Cruz Mountains 450 km north. Associates in the coastal strip were 
three pines that are of coastal occurrence today, Monterey pine (Pinus radiata). Island 
pine {P. remorata), and Stanton pine {Pinus muricata var. stantonii). The Seacliff' 
assemblage suggests that temperature along the coastal strip was approximately 13.5°C 
as compared with 15°C today and that rainfall totalled 76 cm in contrast to 38 cm at 

Development of Modern Forest Geography 

The conifer forests in the mountains of southern California and Baja California 
Norte are now perched as discontinuous stands on isolated mountain ranges at altitudes 
generally above 1 500 m. At lower altitudes, semidesert, chaparral and woodland vege- 
tation thrive under warmer, drier climates. The discontinuous montane forests have 
similar dominants, notably Abies concolor, Calocedrus decurrens, Pinus jeffreyi, P. 
ponderosa, P. lambertiana, P. murrayana, Populus hastata {""trichocarpd") and P. tre- 
muloides (see Griffin and Critchfield 1972, Munz 1935, Nelson 1922, Wiggins 1980). 
In addition, many forbs, herbaceous perennials, and shrubs link these montane forests. 

The now-isolated forest taxa once had a more continuous distribution in the Late 
Pliocene-Early Pleistocene when climate was considerably wetter and cooler than at 
present, as demonstrated by the Soboba flora from San Jacinto Valley in interior 
southern California (Axelrod 1967<3) and by the limited Chula Vista flora. 

In the Plio-Pleistocene transition, mountains were still relatively low, probably 
scarcely half their present altitudes which now reach up to 3000-3400 m. As colder, 
wetter climates spread southward, conifer forests shifted from the Sierra Nevada south- 
ward into the Tehachapi-Mount Pinos-Liebre Mountain region and thence through 
the San Gabriel-San Bernardino-San Jacinto Mountains and into the Sierra San Pedro 
Martir in Baja California Norte (see photos in Nelson 1922). As the fault-block moun- 
tains were elevated later to their present heights, the forests were stranded as disjunct 
stands isolated by drier, warmer climates occupied by woodland, chaparral, sage, and 
semi-desert vegetation. As the forests were restricted to higher altitudes, some taxa 
disappeared locally, thus accounting for the present disjunct occurrences in the moun- 
tains of southern California (see maps in Griffin and Critchfield 1972, Little 1971, also 
discussion in Munz 1935). Other disjunct distributions of Sierran montane taxa in the 
mountains of southern California are exemplified by Acer glabrum, Cornus nuttallii, 
Euonymus occidentalis, Phyllodoce breweri, Rhododendron occidentale, Prunus emar- 
ginata, and many forbs, herbaceous perennials and shrubs listed by Munz (1935). 
Greater disjunctions from the southern Sierra Nevada are seen in the distribution of 
Pinus murrayana (San Gabriel, San Bernardino, San Jacinto mountains, and San Pedro 
Martir) and Populus tremuloides (San Bernardino Mountains and San Pedro Martir). 

Evolution and Biogeography of Pinus radiata 

The Chula Vista flora provides additional evidence regarding the probable evo- 
lutionary history of the five living populations of Pinus radiata. Based on their cones 
these populations can be separated into two groups. The populations of P. radiata on 
Cedros Island (var. cedrosensis) and on Guadalupe Island (var. binata) represent one 
group characterized by small symmetrical to slightly asymmetrical cones without prom- 
inent apophyses except for the extreme variation seen in var. binata. By contrast, the 
other group represented by the California populations at Monterey, Aiio Nuevo and 
Cambria possess larger, regularly asymmetrical cones with large, rounded apophyses. 


Table 3. Trees and shrubs in coastal southern CaHfomia with close relatives in equable montane areas of 

California Mexico 

Arbutus menziesii A. xalapensis 

Ceanothus arboreus C. caeruleus 

Cercocarpus traskiae C. mojadensis 

Comarostaphylis diversifoHa C. spp. ( 1 2 or more) 

Myrica californica M. mexicana 

Pinus remorata P. oocarpa 

Pinus torreyana P. oxacana 

Prunus lyonii P. prionophylla 

Vaccinium ovatum V. confertum 

There is considerable similarity between Pinus radiata var. cedrosensis and P. 
remorata Mason which has large populations in Pine Canyon west of Lompoc, on Santa 
Cruz and Santa Rosa islands, and in Baja California near the coast southwest of San 
Vicente. Cones of Cedros Island pine differ from those off. remorata in having some- 
what more prominent apophyses and slightly thicker cone scales. These differences 
seem to reflect the different environments to which these pines are adapted, with the 
cooler, foggier, moister climate in the north favoring the persistence of a more "prim- 
itive type," as exemplified by P. remorata. Cones of P. remorata are similar to those 
of P. oocarpa, though in its northern areas P. oocarpa cones approach those of P. 
radiata var. cedrosensis in having more prominent apophyses (see U.C. Herbarium) 
than in areas to the south. 

The geologic ages of these pines are not presently known, but to judge from their 
relation to Pinus oocarpa and their present disjunct distribution (see Martinez 1948), 
they have probably been in existence since the Middle Miocene. Up to 500 km of right- 
lateral strike-slip movement since the Middle Miocene along the San Andreas fault 
system and related rifting in the Gulf of California is probably in large part responsible 
for the present biogeography (Gastil and Jensky 1973). The fact that Baja California 
was once part of the Mexican mainland is supported by floristic evidence that provides 
additional data with respect to derivation of Monterey pines from ancient members of 
Pinus subsect. Oocarpeae. Geologic evidence indicates that a chain of low coastal 
mountains was once situated west of the San Andreas rift. These hills were probably 
covered with taxa that now have closely allied species in southern California and in 
the mountains of Mexico, chiefly in areas of equable climate (Table 3). 

Today, these allied taxa are now isolated by the broad stretch of the Sonoran Desert 
and adjacent Thorn Forest vegetation. Links are also provided by a number of taxa 
now in Mexico that have allied fossil species in the Miocene Puente and Modelo floras 
of coastal southern California (Axelrod 1977:162), distributed in Clethra, Magnolia, 
Nectandra, Persea, Quercus, Sabal and others. They clearly indicate that a humid, 
equable Miocene route connected these areas. That there was a near-coastal route is 
indicated also by the Mint Canyon (Axelrod 1979:25, 32) and Tehachapi floras (Axelrod 
1939). They demonstrate that by 19 m.y. ago— and certainly earlier— interior southern 
California-Arizona was too semiarid and hot for the taxa noted above. Hence, allies 
of the mesic species now in coastal California or in Mexico shifted into coastal southern 
California via a near-coastal, not an interior route. It was this route that brought pines 
of subsect. Oocarpeae to California. 

At that time, it is inferred that the ancestor of Pinus radiata var. cedrosensis was 
in the mountains, living under a mild, warm temperate climate. Insularity developed 
from opening of the Gulf of California (since 5 m.y.), and submergence of the outer 
continental shelf There the pine has persisted on the higher summits of Cedros Island 
where it is sheltered by a regular summer fog belt at altitudes above 600 m. The smaller 
cones of the Cedros and Guadalupe Island pines seemingly reflect their closer relation 
to P. oocarpa which was not far away in the mountains of Sonora-Sinaloa, to which 


Baja California was joined in the Miocene. Further, their small size may reflect the 
summer rain regime to which they are and were adapted. 

Pinus diegensis, closely allied to P. radiata var. binata, may have been derived 
from var. cedrosensis by adaptation to somewhat greater aridity over the northern part 
of the earlier distribution of the immediate ancestor of var. cedrosensis. This may 
account for the greater variation in symmetry and apophyses development in P. radiata 
var. binata. That P. diegensis (cf. binata) may represent the ancestral form that gave 
rise to the three California populations of Pinus radiata is suggested by its Pliocene 
mainland occurrence, and by its cone variation which is intermediate between var. 
cedrosensis and the type Monterey population. 

Cones of the three California populations o^ Pinus radiata, which are larger, more 
asymmetrical, and have prominent apophyses, presumably reflect adaptation to a pro- 
gressively more extreme mediterranean climate of dry summers. In the suggested evo- 
lutionary sequence— oocarpa-remorata-cedrosensis-binata-Monterey-A Ho Nuevo- 
Cambria—iX is noteworthy that Pinus diegensis approaches the variation of the Mon- 
terey population. This lends further, though still tenuous, support to its probable place 
in the evolution of Monterey pines. 

The fossil samples now available suggest that Pinus radiata populations with cones 
comparable in size, symmetry, and apophyses development to the Guadalupe and 
Monterey populations were rather widespread in coastal California well into the Pleis- 
tocene. At Carpinteria cones more nearly approaching the larger-sized Aiio Nuevo 
population appear only late in the Pleistocene (Axelrod 1980^). Cones the size of the 
present Cambria population are not now known as fossils. They may be the most recent 
of the group, possibly originating in post-glacial time. 


In the Late Pliocene (ca. 3 m.y.), increased precipitation and lower temperatures 
in California enabled floras of more mesic, cooler requirements to replace floras 
adapted to drier, warmer climate, as illustrated by the Mulholland (Axelrod \944a), 
Oakdale (Axelrod 1 944b), Piru Gorge (Axelrod 1 9506^) and Mount Eden (Axelrod 1937, 
\950b) floras. The more mesic, younger floras, like the Santa Rosa, Napa, Upper 
Wildcat, Turlock and (probably) Coalinga, still have a few exotic taxa allied to species 
in summer rain areas, reflecting a warmer sea surface than that of today. Owing to its 
more southerly position, the influence of cooler climate at Chula Vista was not so 
pronounced, though precipitation was about double that of today. The lowland flora 
of southern California then represented a different fforistic province than that to the 
north, one adapted to a warmer climate, much as comparable thermal differences 
separate the floras of northern and southern California today. 

The succeeding colder, wetter climate of the latest Pliocene and early Pleistocene 
enabled Douglas fir {Pseudotsuga menziesii) forest to shift south into coastal southern 
California, fully 350 km (or somewhat more) south of its present scattered coastal 
stations. Over the interior, lowered temperature and higher precipitation enabled the 
Sierran mixed conifer forest to range southward into the mountains of southern Cal- 
ifornia and Baja California Norte. The Sierran mixed conifer forest then lived fully 
1000 m below its present level at a time when mountains were appreciably lower. The 
present forests were stranded as isolated patches as the discontinuous mountains were 
uplifted in the middle and later Quaternary and drier, hotter climates spread over the 
region. These later, more extreme conditions probably account for the present disjunct 
occurrences of diverse montane taxa in the mountains of southern and Baja California, 
and for the restriction of taxa allied to those in the Chula Vista flora to more local 

Evidence suggests that Pinus diegensis (cf. P. radiata var. binata) may have given 
rise to the California populations of Pinus radiata by progressive adaptation to in- 
creasing summer drought. Further, P. radiata var. binata did not originate in insular 


isolation, but like many other insular endemic trees and shrubs, was confined there by 
unfavorable land climates during the past. 


Family Pinaceae 

Pinus diegensis new species 

Figures 6A, B, 7A, B 

Description.— Cones long-oval to elliptic oval; small cone (SDSNH 25135) 9 cm 
long and 5 cm broad, medium sized cone (SDSNH 25 165) about 10-1 1 cm long(estim.), 
and about 6.5 cm broad; largest cones (SDSNH 25136, 25137) somewhat flattened, 
fragmentary. Cone scales up to 4.0-4.5 cm long and 2.0-2.3 cm broad, distally. Tips 
of cone scales broadly flattened, with slightly swollen, convexly rounded apophyses 
well inside tip of scale. 

Types. -Holotype SDSNH no. 25135, Paratypes SDSNH no. 251 10, 25136, 25137, 

Discussion. — Four incomplete cones are sufliciently distinct from those previously 
described to warrant the recognition of a new species. Examination of the large, excellent 
collection of pine cones in the herbarium of the U.S. Forest Service, Institute of Ge- 
netics, Placerville, California, indicates that P. diegensis is a member of the subsect. 
Oocarpeae (Little and Critchfield 1 969). The fossil cones are most similar to the variable 
cones produced by Pinus radiata var. binata Engelmann from Guadalupe Island. The 
variation present in the small fossil collection is readily duplicated by the large suites 
of cones that have been recovered from the island. 

Howell (1941a) discussed cone variation in the Guadalupe pine, noting that asym- 
metrical cones with large apophyses on the outer side (like fig. 6B) were similar to the 
type which comes from Monterey. A second form with nearly symmetrical cones (like 
fig. 6A) and with scales alike on all sides and with little prominent apophyses devel- 
opment Howell termed forma guadalupensis. A third form, intermediate between the 
latter two he termed /orma binata, noting that it corresponded with the type specimen 
collected from Guadalupe. As discussed separately in the section on Evolution, the 
variable binata on Guadalupe Island may represent a population close to that which 
gave rise to the three living populations in central California. 

In southern California, fossil pine cones allied to the present California populations 
of Pinus radiata have been recorded previously from the late Tertiary Mount Eden 
(Axelrod 1937) and Lower Pico floras (Dorf 1930). It is also known from Pleistocene 
floras at Carpinteria (Chaney and Mason 1933), near Seacliff on the coast west of 
Ventura (Axelrod 1983), and at Rancho La Brea (Mason 1927). In coast-central Cal- 
ifornia, it is known from Mussel Rock on the outer coast south of San Francisco. Here 
it has been recovered from an old forest soil that rests on Franciscan diabase and is 
overlain by the basal beds of the marine Merced Formation, approximately 5-6 m.y. 
old (Axelrod 1967Z>). Pleistocene records in central California are near Pt. Sal (Axelrod 
1961b), Little Sur (Langenheim and Durham 1963), Millerton (Mason 1934, Axelrod 
1980a), and Drakes Bay (Axelrod 1980a, 1983). 

The present record is the most southern fossil locality now known for species of 
Monterey pine. 

Pinus jeffreyoides new species 
Figures 7C, D 

Description.— Cone large, estimated 15-10 cm long and fully 10 cm broad; cone 
scales large, up to 4 cm long in central part of cone; terminal part of cone scale broadly 
triangular, 1.5-2.5 cm wide and 0.8-1.1 cm thick as measured below the umbo. 

Typ^'^.-Holotype SDSNH 25166; Paratypes SDSNH 25138. 25167. 

Discussion.— This is a member of subsect. Ponderosae Little and Critchfield ( 1 969) 
and is well-matched by cones of the living Pinus jeffreyi Greville and Balfour. Rela- 



J jtff^ 

Figure 6. \,^—Pinus diegensis new species, A — holotype, SDSNH 25135, B— paratype, SDSNH 25165. 
All specimens x ] (latex casts). 

tionship with P. jeffreyi is apparent in the morphology of the distal ends of the cone 
scales, which are much thicker than those of the allied P. ponderosa Lawson. 

Pinus pieperi Dorf 
Figures 7B 

Pinus pieperi Dorf, Carnegie Inst. Wash. Publ. 412, p. 69, pi. 5, figs. 7-10, 1930; 
Axelrod, Carnegie Inst. Wash. Publ. 476, p. 156, pi. 2, figs. 2, 3, 1937. 

Referred specimen. -S,X)SN\\ 25168. 

Remarks. — A. single fragment of a cone with distinctive cone scales that have 


Figure 7. A,B—Pinus diegensis new species. A— paratype, SDSNH 251 10, B— paratype, SDSNH 25137; 
CD- Pinns jeffreyoides new species, C-holotype, SDSNH 25166, D-paratype, SDSNH 25138; E-Pinus 
pieperi Dorf, SDSNH 25168. All specimens x 1 (latex casts). 

prominently hooked, large apophyses represents this species which is allied to digger 
pine, Pinus sabiniana Douglas of central California. The specimen measures 6.0 cm 
long and 5.5 cm wide. 

Pinus pieperi has been recorded previously in southern California from the Plio- 
Pleistocene rocks north of Ventura (Wiggins 1951). in the lower part of the Pico For- 
mation west of Ventura (Dorf 1930). in the upper Pico Formation near Seacliff on the 
coast west of Ventura (Axelrod 1 983). as well as in the Mount Eden flora near Beaumont 
(Axelrod 1937). A fragmentary cone scale from the Anaverde Formation near Palmdale 


has also been referred to Pinus pieperi (Axelrod 1950c). Digger pine may have been 
eliminated from southern California as drier hotter climate spread there during the 

Family Arecaceae 

Gen. et sp. indet. 

Figure 8A 

Referred specimen.— SUSNH 25 163. 

Remarks.— A fragmentary specimen that certainly represents a palm is in the 
collection. It is 7 cm long and 2.5 cm broad, the blade has 4 rays each with a prominent 
midvein and each is bordered by 7-8 fine parallel veins that are somewhat less than 
1 mm apart. 

Comparison with several genera indicates that it may represent a species of Sabal 
or Brahea {=Erythea), but in view of the incomplete nature of the fossil reference to 
either (or any) genus seems unjustified. Both of the noted genera extend up into oak 
woodland vegetation, Sabal in northern Sinaloa and adjacent Sonora, Brahea in the 
highlands of southernmost Baja California, and both regions with ample rainfall in 
summer. In addition, Brahea is associated with Monterey pine on Guadalupe Island. 

Family Juncaceae 

Gen. et. sp. indet. 

Figure 83 

Referred specimen. -SDSNH 25164. 

Remarks.— A single slender reed-like leaf impression is 5 cm long and 7 mm wide 
with 9-10 parallel veins. The blade is without a midrib and clearly represents a sedge- 
or reed-like plant of indeterminate nature. It seems comparable to some leaves of 
Cyperus, and especially to Juncus, both common in marshy areas and on banks along 
slow, meandering rivers. 

Family Salicaceae 

Populus alexanderi Dorf 

Figures 8E, F 

Populus alexanderi Dorf, Carnegie Inst. Wash. Publ. 412, p. 75, pi. 6, fig. 11 only, 
1930; Axelrod, Carnegie Inst. Wash. Publ. 553, p. 281, pi. 48, fig. 4, 1944; Axelrod, 
Univ. Cahf Publ. Geol. Sci. 34, p. 128, pi. 19, figs. 1-11, 1958. 

Referred specimens. -^USHH 25159, 25160, 25161, 25162. 

Remarks.— This fossil species represents a cottonwood similar to the coastal Pop- 
ulus trichocarpa Torrey and Gray, the type of which comes from the Santa Clara River 
near Ventura. This species differs from leaves commonly identified as P. alexanderi 
and P. eotremuloides Knowlton in its ovate shape and smaller size as compared with 
the lanceolate-ovate form of the species from Miocene floras to the north and in the 
north coast Pliocene floras as well. The specimens from the Verdi flora (Axelrod 1958, 
pi. 19) illustrate the nature of the species, and the accompanying plate 20 (loc. cit.) 
shows how much Populus ""trichocarpa''^ from the mountains differs from it; this mon- 
tane and north coastal species seems to represent P. hastata Dode. 

P. alexanderi (restricted) is indicative of a climate with mild winters as compared 
with P. eotremuloides which inhabited areas with colder climate. The ovate-leaved P. 
alexanderi is related to P. emersoni Condit from the San Pablo flora (Condit 1938; 
Lesquereux 1883, pi. 55, figs. 3, 5, only), though the latter has consistently larger leaves. 

Salix wildcatensis Axelrod 
Figures 8C, D 

Salix wildcatensis Axelrod, Carnegie Inst. Wash. Publ. 553, p. 132, 1944 (see synon- 
ymy); Chaney, Carnegie Inst. Wash. Publ. 553, p. 341, pi. 58, Fig. 2, 1944. 


Figure 8. A-Arecaceae indet., SDSNH 25163; B-Juncaceae indet., SDSNH 25164; C,D-Salix wild- 
catensis Axelrod, C-SDSNH 25 1 53, D-SDSNH 25 1 52; E,F - Populus alexanderi Dorf, E-SDSNH 25 1 59, 
F-SDSNH 25160. All specimens x 1. 


Salix coalingensis Dorf (in part), Carnegie Inst. Wash. Publ. 476, p. 170, pi. 4, fig. 8, 

Referred sped mens. -SDS^H 25 1 52, 25 1 53, 25 1 54. 

Remarks. — Three fragmentary specimens, the largest 7.0 cm long and 2.6 pm broad, 
represent this willow. It is allied to Salix lasiolepis Bentham, a common shrub to small 
tree that is widely distributed in the Coast Ranges and lower Sierra Nevada, reaching 
southward into the coastal slopes of northern Baja California. It is also disjunct to 
southeastern Arizona. In both areas it is a common member of riparian vegetation in 
oak woodlands. 

Family Fagaceae 

Quercus lakevillensis Dorf 

Figure 9C 

Quercus lakevillensis Dorf, Carnegie Inst. Wash. Publ. 412, p. 82, pi. 8, figs. 4, 5, 1930; 
Axelrod, Carnegie Inst. Wash. Publ. 590, p. 58, pi. 3, fig. 4, 1950; Axelrod, Univ. 
Calif. Publ. Geol. Sci. 121, p. 165, pi. 18, figs. 2, 3, 1980. 

Referred specimen. -SI>S^H 25158. 

Remarks.— A single leaf in the flora is similar to those produced by the common 
California live oak, Quercus agrifolia Nee. The specimen is oval in outline, has broad 
sinuses and wavering, irregular secondaries that diverge at moderate angles. The modem 
species is common in the Coast Ranges from Sonoma County southward into Baja 
California. Near the coast, where it is subject to regular summer fog, it forms pure 
dense woodlands. Elsewhere it is a member of diverse communities, including redwood 
forest, closed cone pine forest, broadleaved sclerophyll woodland and occurs also in 
the coastal sectors of digger pine woodland. 

Family Lauraceae 

Persea coalingensis (Dorf) Axelrod 

Figures 9A, B 

Persea coalingensis (Dorf) Axelrod, Carnegie Inst. Wash. Publ. 553, p. 132, 1944 (see 
synonymy and discussion); Axelrod, Univ. CaHf Publ. 121, p. 112, pi. 12, fig. 
4; p. 167, pi. 19, fig. 7; pi. 20, figs. 2-4, 1980. 

Referred specimens. -^US>N¥[ 25149, 25150, 25151. 

Remarks.— The long-elliptic leaves of avocado are the commonest leaf fossil in 
the Chula Vista flora. Most were broken during transport into the marine basin though 
sufficient details of venation are present to permit their certain reference to this late 
Tertiary species. Two nearly complete specimens tentatively grouped under Persea may 
represent Magnolia but the finer details of venation which would make their identifi- 
cation certain are not preserved. 

Family Platanaceae 

Platanus paucidentata Dorf 

Figures lOD, E 

Platanus paucidentata Dorf, Carnegie Inst. Wash. Publ. 412, p. 94, pi. 10, figs. 4, 9; 
pi. 11, fig. 1; pi. 12, fig. 1, 1930; Axelrod, Carnegie Inst. Wash. Publ. 476, p. 174, 
pi. 5, figs. 4, 5, 1937; Axelrod, Univ. Calif. Publ. Geol. Sci. 121, p. 113, pi. 13, 
fig. 1; p. 168, pi. 20, fig. 1, pi. 21, fig. 7, 1980. 

Referred specimens. -S\)S\<^H 25155, 25156, 25157. 

Remarks.— The typical lobed leaves of this sycamore are in the flora. The large 
leaves regularly produced by the species were broken and mangled during transport 
into the marine basin. However, the distinctive primary venation and the lobed nature 
of the leaves permits their ready identification. 


Figure 9. A,B-Persea coalingensis (DorO Axelrod, A-SDSNH 25150, B-SDSNH 25151; C-Quercus 
lakcvillcnsis DorfSDSNH 25158; D,E- Platanus paucidentataDorf , D-SDSNH 25155, E-SDSNH 25157. 
All specimens x 1. 


The fossil species has leaves similar to those of Platanus racemosa Nuttall, found 
along stream banks and floodplains in oak woodland vegetation from central California 
southward into Baja California. An allied species. P. wrightii Watson, is in Arizona 
and border areas. The fossil species, P. paucidentata, is common in the Miocene and 
Pliocene floras of California. * 


The cooperation of Watt Industries, Rancho Santa Fe, California, and Financial 
Scene, San Diego, California, is gratefully acknowledged for permitting and supporting 
the collection of fossils on their Chula Vista property. 

In addition the Planning Department, City of Chula Vista and especially Douglas 
Reid of that office are to be commended for realizing the significance of paleontological 
resources within their jurisdiction. 

Special thanks are extended to Richard A. Cerutti and Bradford O. Riney who 
collected most of the paleobotanical material discussed herein. 

The junior author's work on this project was supported in part by grants from the 
Parker Foundation, the Scripps Foundation, and the J. F. Sefton Foundation. The 
additional support of Joseph and Joanne Parker of Comado, California, is also gratefully 

The senior author worked on the project during tenure of National Science Foun- 
dation Grant DEB 80-25525, which is acknowledged with thanks. 

Literature Cited 

Axelrod, D. I. 1937. A Pliocene flora from the 
Mount Eden beds, southern California. Car- 
negie Institution of Washington Publication 

. 1939. A Miocene flora from the western 

border of the Mohave Desert. Carnegie Insti- 
tution of Washington Publication 516:129. 

. 1944a. The MulhoUand flora. Carnegie 

Institution of Washington Pubhcation 553:103- 

. 1944/?. The Oakdale flora. Carnegie In- 
stitute of Washington Pubhcation 553:147-165. 

. 1944r. The Sonoma flora. Carnegie In- 
stitute of Washington Publication 553:167-206. 

. 1 950(2. A Sonoma florule from Napa, Cal- 
ifornia. Carnegie Institute of Washington Pub- 
lication 590:23-71. 

. 1950^. Further studies ofthe Mount Eden 

flora, southern California. Carnegie Institute of 
Washington Publication 590b:73-l 17. 

. 1950c. The Anaverde flora of southern 

California. Carnegie Institute of Washington 
Publication 590:119-158. 

. \950d. The Piru Gorge Flora of southern 

California. Carnegie Institute of Washington 
Publication 590:159-214. 

— . 1958. The Pliocene Verdi flora of western 
Nevada. University of California Publications 
in Geological Sciences 34:91-160. 

— . 1967a. The Pleistocene Soboba flora of 
southern California. University of California 
Publications in Geological Sciences 60:1-108. 

— . 1967^. Evolution of the Califomian 
closed-cone pine forest. Pp. 93-150 in R. N. 
Philbrick, (ed.). Proceedings ofthe Symposium 
on Biology ofthe California Islands. Santa Bar- 
bara Botanical Garden. 

— . 1976. History ofthe coniferous forest. 

California and Nevada. University of Califor- 
nia Publications in Geological Science 70:1- 

— . 1977. Outline history of California vege- 
tation. Pp. 139-193 in M. G. Barbour and J. 
Major (eds.). Terrestrial Vegetation of Califor- 
nia. J. Wiley & Sons, New York. 

— . 1979. Age and Origin of Sonoran Desert 
Vegetation. California Academy of Sciences 
Occasional Papers 132:1-74. 

— . 1980a. History of the maritime closed- 
cone pines. Aha and Baja California. Univer- 
sity of California Publications in Geological 
Sciences 120:1-143. 

— . 1 980^. Contributions to the Neogene Pa- 
leobotany of Central California. University of 
California Publications in Geological Sciences 

1983. New Pleistocene conifer records, 

coastal California. University of California 
Publications in Geological Sciences 127:1-107. 

Bailey, H. P. 1960. A method of determining the 
warmth and temperateness of climate. Geo- 
grafiska Annaler 42:1-16. 

. 1964. Toward a unified concept ofthe 

temperate climate. Geographical Review 54: 

Barnes, L. G. 1 973. Pliocene cetaceans ofthe San 
Diego Formation. Pp. 37-43 in Arnold Ross 
and R. J. Dowlen (eds.). Studies on the geology 
and geological hazards ofthe greater San Diego 
area, California. San Diego Association of Ge- 

. 1976. Outline ofthe eastern North Pacific 

fossil cetacean assemblages. Systematic Zool- 
ogy 25(4):32 1-343. 

Chaney, R. W., and H. L. Mason. 1933. A Pleis- 
tocene flora from the asphalt deposits at Car- 


pinteria, California. Carnegie Institute of 
Washington Publication 415:45-79. 

Condit, C. 1938. The San Pablo Rora of West 
Central California. Carnegie Institute of Wash- 
ington Publication 476:217-268. 

Demere, T. A. 1983. The Neogene San Diego 
Basin: A review of the marine Pliocene San 
Diego Formation. Pp. 187-195 in D. K. Larue 
and R. J. Steel (eds.). Cenozoic marine sedi- 
mentation Pacific Margin, U.S.A. Society of 
Economic Paleontologists and Mineralogists, 
Pacific Section. 

Dorf, E. 1930. Pliocene floras of California. Car- 
negie Institute of Washington Publication 412: 

Gastil, R. G., and W. Jensky. 1973. Evidence of 
strike-slip displacement beneath the trans- 
Mexican volcanic belt. Stanford University 
Publications in Geological Sciences 12:171- 

Gentry. H.S. 1942. Rio Mayo Plants: A study of 
the flora and vegetation of the Valley of the 
Rio Mayo, Sonora. Carnegie Institute of Wash- 
ington Publication 527:1-328. 

. 1946. Notes on the vegetation of Sierra 

Surotato in northern Sinaloa. Torrey Botanti- 
cal Club Bulletin 73:451-462. 

Grant, U. S., IV, and H. R. Gale. 1931. Catalogue 
of the marine Pliocene and Pleistocene Mol- 
lusca of California and adjacent regions. San 
Diego Society Natural History Memoir 1:1- 

Griffin, J. R., and W. B. Critchfield. 1972. The 
distribution of forest trees in California. USDA, 
Forest Service Research Paper PSW-82/1972. 
118 pp. 

Hertlein, L. G., and U. S. Grant, IV. 1944. The 
geology and paleontology of the marine Plio- 
cene of San Diego, California, Pt. I, Geology. 
San Diego Society Natural History Memoir 2: 

. 1 954. Geology of the Oceanside-San Die- 
go coastal area, southern California. Pp. 53- 
63 in R. H. Jahns (ed.). Geology of Southern 
California, Chap. II, Geology of the Natural 
Provinces. California Division of Mines Bul- 
letin 170. 

. 1960. The geology and paleontology of 

the marine Pliocene of San Diego, California, 
Pt. 2a, Paleontology. San Diego Society Nat- 
ural History Memoir 2a:73-I33. 

1972. The geology and paleontology of 

the marine Pliocene of San Diego, California, 
Pt. 2b, Paleontology. San Diego Society Nat- 
ural History Memoir 2b: 143-409. 

Howard, H. 1 949. New avian records for the Plio- 
cene of California. Carnegie Institute of Wash- 
ington Publication 584:177-199. 

Howell, J. T. 1941a. The closed-cone pines of 
insular California. Leaflets of Western Botany 

. 1941/7. My visits to Guadalupe Island. 

Leaflets of Western Botany 3:36-41. 

Ingle, J. C, Jr. 1967. Foraminiferal biofacies, 
variations and the Miocene-Pliocene bound- 
ary in southern California. Bulletin American 
Paleontology 52(236):2 16-394. 

Keen, A. M., and H. Bentson. 1944. Checklist of 

California Tertiary marine Mollusca. Geolog- 
ical Society of America Special Papers 56:1- 

Kennedy, M. P. 1975. Geology of the San Diego 
Metropolitan area, California, Section A. Cal- 
ifornia Division Mines and Geology Bulletin 

Lajoie, K. R., A. M. Sama-Wojcicki, and R. F. 
Yerkes. 1982. Quaternary chronology and 
rates of crustal deformation in the Ventura area, 
California. Neotectonics of the Ventura Basin. 
Field Trip No. 3:43-51. Guidebook for 78th 
Ann. Meeting Cordilleran Section, Geological 
Society of America, Anaheim, California. 

Langenheim, J. H., and J. W. Durham. 1963. 
Quaternary closed-cone pine forest from trav- 
ertine near Little Sur, California. Madrono 1 7: 

Lesquereux, L. 1883. Contributions to the fossil 
flora of the Western Territories. III. The Cre- 
taceous and Tertiary floras. U.S. Geological 
Survey of Territories, vol. 8:1-283. 

Libby, W. J., M. H. Bannister, and Y. B. Linhart. 
1968. The pines of Cedros and Guadalupe 
Islands. Journal of Forestry 66:846-853. 

Little, E.L., Jr. 1971. Atlas of United States Trees. 
U.S. Department of Agriculture Forest Service, 
Miscellaneous Publication 1 146. 200 maps. 

, Jr., and W. B. Critchfield. 1969. Subdi- 
visions of the genus Pinus (Pines). U.S. De- 
partment of Agriculture Forest Service, Mis- 
cellaneous Publication 1 144:1-51. 

Mandel, D. J., Jr. 1973. Latest Pliocene Fora- 
minifera in the upper part of the San Diego 
Formation, California. Pp. 33-36 in A. Ross 
and R. J. Dowlen (eds.). Studies on the geology 
and geologic hazards of the greater San Diego 
area, California. San Diego Association of Ge- 

Martinez, M. 1948. Los Pinos Mexicanos. 2nd 
ed. Mexico: Ediciones Botas. 316 pp. 

Mason, H. L. 1927. Fossil records of some west 
American conifers. Carnegie Institute of Wash- 
ington Publication 346:139-158. 

. 1934. Pleistocene flora of the Tomales 

Formation. Carnegie Institute of Washington 
Publication 415:81-179. 

Miller, L. 1 956. A collection of bird remains from 
the Pliocene of San Diego, California. Califor- 
nia Academy of Sciences Proceedings 28( 1 6): 

Munz, P. A. 1935. A Manual of Southern Cali- 
fornia Botany. Claremont, California, Clare- 
mont College. 

Nelson, E. W. 1922. Lower California and its Nat- 
ural Resources. National Academy of Sciences 
Memoir 16:1-194. 

U.S. Department of Agriculture, Weather Bureau. 
1938. Atlas of Climatic Charts of the Oceans. 
W.B. No. 1247. 130 charts. 

U.S. Navy. 1956-58. Climatic Atlas of the World. 
Washington, D.C. 

Vedder, J. G. 1 972. Review of stratigraphic names 
and megafaunal correlation of Pliocene rocks 
along the southeast margin of the Los Angeles 
basin, California. Pp. 158-172, in E. H. Stine- 
meyer (ed.). Pacific Coast Miocene Biostrati- 


graphic Symposium. Society of Economic Pa- . 1980. Flora of Baja California. Stanford: 

leontologists and Mineralogists. Standford Univ. Press. 1025 pp. 

Wiggins, I. L. 1951. An additional specimen of Woodring, W. P., and M. N. Bramlette. 1951. Ge- 

Pinus pieperi Dorf from Ventura County, Cal- ology and paleontology of the Santa Maria dis- 

ifomia. American Journal of Botany 38:21 1- trict, California. U.S. Geological Survey 

213. Professional Paper 222:1-185. •■ 




Volume 20 Number 16 pp. 301-312 20 November 1984 

Relationships within Eumalacostracan Crustacea ^p 

Frederick R. Schram 

San Diego Natural History Museum, P.O. Box 1390, San Diego, CA 92112 USA 


Abstract. A cladistic analysis was performed on 20 constituent higher taxa within the Eumala- 
costraca based on 3 1 characters of external anatomy. Variants of the most parsimonious scheme are 
presented, and the effects of tolerating different levels of uncertainty are evaluated. It is concluded that: 
1) while the basic outline of Caiman's (1904) taxonomy of Eumalacostraca might be utilized, the 
arrangement within peracarids postulated by Siewing (1956) cannot be maintained; 2) the Baupldne 
approach of Schram (1981) has some merit and some of the controversial higher taxonomic groupings 
of eumalacostracan "orders" originally indicated by that method are vindicated; 3) the idea that the 
carapace is a derived feature within eumalacostracans, advanced by Dahl (1983), can be maintained 
only if a high level of homoplasy is tolerated; 4) the concept of a taxon Mysidacea seems best abandoned. 


The basic modem classification of eumalacostracan crustaceans was outlined by 
Caiman (1904, 1909) with little reference at that time to what the details of phyletic 
relationships between and within groups might have been. However, it was Siewing 
(195 1, 1956) who presented a phylogenetic tree for eumalacostracans widely subscribed 
to by subsequent authorities (e.g.. Fryer 1964, Hessler 1969). 

Recently, however, the Calman/Siewing scheme for Eumalacostraca sensu sthcto 
has been questioned. Schram (1981) recognized basic structural plans within the Eu- 
malacostraca, but the methodology he employed was limited by the number of char- 
acters that could be handled essentially by pencil and paper. However, the method was 
helpful in three respects. First, it illustrated a variable range of possible dendrograms. 
Each variant dendrogram was constrained by which characters received initial emphasis 
and, thus, demonstrated the basic range of uncertainty that must be implicit in any 
phylogenetic analysis. Second, the analysis suggested certain "supraordinal" relation- 
ships which were a bit unexpected, especially within the peracarid groups. For example, 
isopods and amphipods were united; and cumaceans, tanaids, and spelaeogriphaceans 
were allied to each other with some suggestion of more distant possible links of these 
to thermosbaenaceans. Third, the method also produced a number of "paper" Baupldne 
which were not occupied or had yet to be discovered. Implicit in these hypothetical 
morphotypes was the idea that if the method had any merit at all, some of those 
"empty" Baupldne might eventually be found. 

Watling (1981, 1983) questioned the unity of the superorder Peracarida as a natural 
taxon. He produced two different cladograms for the peracarids. His stated purpose (in 
Schram, 1983:347) was to search for ". . . Baupldne that include the fine structure . . . ," 
and he believed that ". . . the first step in the analysis is to look at all these structures 
for pattern . . . ." In this respect Watling (1983) performed a useful function by focusing 
attention on characters that had largely been overlooked by previous workers such as 
mandible function, maxillipede form, and patterns of arterial circulation. 

Dahl (1983) formally proposed a concept that had been implicit in several of his 
earlier papers, viz., that the lack of a carapace is a primitive feature, that the evolution 
of the carapace had occurred independently several times, and that Caiman's caridoid 


facies was a set of convergent phenomena. Dahl presented some interesting observations 
on comparative carapace development related to these ideas. Watling (especially 1983) 
acknowledged his indebtedness to Dahl's concept of the carapace as a derived feature. 

Finally, Hessler (1983) produced a "defense" of the caridoid facies in which he 
attempted cladistic analysis of the Siewing scheme for peracarids in a mpre formal 
manner than had ever been presented before. Hessler's study, however, produced a 
scheme in which the Siewing arrangement of taxa could be retained only by tolerating 
a great deal of convergence (10 of his 23 characters are convergent in whole or part to 
one or more of the others). 

Thus, several items bear on the problem of eumalacostracan relationships and 
demand some sort of a resolution. First, is the need to assess relationships among 
eumalacostracans by the use of as many characters as possible, and use these characters 
across-the-board for all taxa, fosssil and Recent. Second, a test is demanded both for 
Dahl's concept of the carapace as a derived feature, as well as some of the "strange" 
higher groupings suggested by Schram (1981). And third, it is necessary to arrive at a 
scheme which will group the taxa in question strictly by their shared derived character 
states with the fewest number of convergences possible. 


One way to analyze large numbers of characters and taxa so as to achieve the most 
parsimonious arrangement, based solely on shared derived characters, is to use one of 
the various versions available for the Wagner 78 program. For this analysis, I decided 
to "break up" certain large and diverse traditional eumalacostracan taxa and treat their 
components as separate units to test both the viability of such taxa and the "reason- 
ableness" of the characters used. To this end the suborders of Mysidacea (Mysida, 
Lophogastrida, and Pygocephalomorpha) and of Decapoda as outlined by Burkenroad 
(1981) (Dendrobranchiata, Eukyphida, Euzygida, and Reptantia) were evaluated as 
separate entities. The choice of taxa for the decapods was somewhat arbitrary since, 
for example, Felgenhauer and Abele (1983) break the Eukyphida into two groups 
coequal with the others, Procarididea and Caridea. 

The 3 1 characters used for this analysis were selected by repeated trial and error 
(as is standard in any computer-generated cladistic treatment of such data), rejecting 
potentially useful characters which had low consistency indexes (i.e., high homoplasy 
values). The ultimate aim of these initial assessments of potentially useful characters 
was to yield the most parsimonious and congruent cladogram possible. Only characters 
that could be assessed for all groups relatively unambiguously were used. For example, 
I did not use the lacinia mobilis because I do not feel its homology has been dem- 
onstrated. As has been shown recently (Dahl and Hessler, 1982), this character is not 
only present in several groups, but is developed differently in these taxa. How can one 
compare the massive laciniae of lophogastrids with the rather delicate ones in other 
peracarids? Or, how are larval laciniae to be judged in comparison to those of adults? 
More needs to be known about the development and functional morphology of laciniae 
before they can be adequately assessed in a phylogenetic analysis. Other characters 
were not used because, while they serve to characterize specific taxa, they are quite 
homoplastic and are known to occur convergently in widely separated groups. For 
example, the use of the presence of second or third maxillipedes was avoided in the 
final analysis since it only served to confirm groupings achieved more effectively by 
singularly derived features. The characters eventually settled upon are given in Table 
1, the numbers indicated corresponding to those used in the cladograms. 

The program was run using several different outgroups, Hoplocarida, leptostracan 
Phyllocarida, and a hypothetical ancestor arbitrarily designated primitive for all 31 
characters. No differences in any of the resultant eumalacostracan cladograms were 
noted. Among other parameters, the program also calculated total lengths of trees (i.e., 
the total number of incidences of derived characters in the cladogram) and the total 
homoplasy value (i.e., a measure of the total array of convergences and character 


Table 1 . Opposing list of character states used in the analysis of relationships within Eumalacostraca. 
Numbers correspond to those used in cladograms. 



1 . Non-caridoid musculature 

2. No zoeal larvae 

3. Carapace not fused to all thoracomeres 

4. No petasma 

5. First thoracomere free of head 

6. Maxillipede with epipodite 

7. No brood pouch formed by first pleopod 

8. No scaphognathite 

9. First thoracopod unmodified 

10. Maxillipede endopod robust 

1 1 . Eggs not brooded on pleopods 

12. No caridean lobe 

13. Biramous thoracopods 

14. All pleopods present 

15. First thoracopod unmodified 

16. Pereiopodal epipodite gills 

17. First thoracopod unmodified 

18. No marsupium 

19. Thoracic endopods non-filtratory 

20. No male cones 

2 1 . Thoracic coxae unmodified 

22. Eyes stalked or lobed 

23. One pair of uropods 

24. Pleopods non-respiratory 

25. Carapace not short 

26. Eggs not brooded under carapace 

27. Maxillipedal epipodite if present simple 

28. Rostrum simple 

29. Thoracic exopods non-respiratory 

30. Maxillipedal epipodite as a single segment 

31. Carapace 

Character reversal used in analysis portrayed in Figure 3 

3 1 . No carapace carapace 

caridoid musculature 


carapace fused to all thoracomeres 


first thoracomere fused to head 

maxillipede without epipodite 

brood pouch between first pleopod and 
venter of thorax 

scaphognathite on maxilla 

maxillipedes with lamellate protopod, 
coxal/basal endites directed mediad 

maxillipede endopod flagelliform 

eggs brooded on pleopods 

caridean lobe on maxillipede 

uniramous thoracopods 

pleopods lost or reduced 

maxillipedes with tendency to form 
gnathobasic endites, endopod pediform 

no pereiopodal epipodite gills 

maxillipedes with basal endites lobate 
and directed distad 

oostegite marsupium 

thoracic endopods filtratory 

male cones 

thoracic coxal plates 

eyes sessile 

more than one pair of uropods 

pleopods respiratory 

carapace short 

eggs brooded under carapace 

epipodite specialized as cup- or spoon- 
like respiratory organ 

pseudorostrum and maxillipedal siphons 

thoracic exopods respiratory 

epipodite with tendency to form as 2-3 

carapace absent 

reversals in the cladogram). These factors proved useful in qualitatively comparing 
different cladograms. 


The computer program generated several variant cladograms. That variant which 
was most parsimonious and yielded the fewest number of convergences and character 
reversals is given in Figure 1. In the series of cladograms summarized in Figures 1-3, 
previous outgroup analysis indicated that the presence of a carapace should be treated 
as primitive because it is present in all hoplocaridans and phyllocaridans. As can be 
seen, the program produced (Fig. 1) an unresolved polychotomy with four branches at 
the base of the Eumalacostraca: eucarids, belotelsonids, syncarids, and waterstonellid/ 
peracarids. A variant of this scheme (Fig. 2) yields an unresolved polychotomy of five 
branches. Although the latter cladogram has the same number of convergences as the 
former, it is somewhat shorter than that of Figure 1 . A convergence in the secondary 
reevolution of pereiopodal epipodite gills between Mysida and Amphipoda is traded 
off for a convergence in the primary loss of pereiopodal epipodite gills in Watersto- 




•^> ^* -^^ ^'^ ;^ >" / ^^ / r / s?' / / / * / 
/ ./ / / / € / / / / / ■/ / / / / / / # ^^ 

<^ ^? ^? / / sf -^ «* ■r"' *' ** *-■ <^ -/■ ^ / ^ i? ^-^ ^ 




Figure 1 . Cladistic relationships of component taxa of the Eumalacostraca, the presence of a carapace 
considered primitive. This is the shortest cladogram with the lowest homoplasy value, the base of the 
cladogram with an unresolved quadrochotomy between eucarids, belotelsonids, syncarids, and waterstonel- 
lids/peracarids. D primitive,  derived. 

nellidea and the "peracarid" line above Mysida. Wagner 78 is designed to produce the 
best resolved cladogram possible from the data given and, thus, the preferred version 
is that seen in Fig. 1 . If on the other hand we wish to tolerate a slightly greater degreee 
of uncertainty (Schram, 1983), then we may choose the variant of Fig. 2 in which 
peracarids can be recognized as a distinct lineage. The relationships indicated in Figure 
1 , however, are not without considerable biological interest. The thrust of the early 
evolution of the waterstonellid/peracarid line was towards increasing specialization of 
thoracopods. First the primitive respiratory epipodites were lost, then oostegites and 
maxillipedes were evolved, and finally some further specializations occurred in specific 
lineages such as filtratory endopods in mysidans (Attramadal, pers. comm.), and further 
maxillipedal and ambulatory modifications in pygocephalomorphs (Schram, 1974). 


>i» A •> -if 4^ .-s? ^ i«^ i^^ 



»* # / ./ / / «» / / / .. i ^. ^^ # ./ / / ^«' 


^^ sf sf <f / <v^ ^^ ^^ t" <^ ^'^ ^ -^ -^ -^ ^"^ ^"^ -^ ^-^ ^ 

^".^"/^<s"'.f ^^ J^^ 

Figure 2. Cladistic relationships of component taxa of the Eumalacostraca, the presence of a carapace 
considered primitive. A variant cladogram from that of Fig. 1 exhibiting ( 1 ) unresolved quintichotomy 
at the base that allows a separation of waterstonellids and peracarids (which would shorten the tree, not 
involve any change in the number of convergences over that of Fig. 1 , but would inject a higher level of 
uncertainty into the cladogram) and (2) an association of pygocephalomorphs as a sister group of the brachy- 
caridans (which would not involve a lengthening of the cladogram but would inject one extra character 
reversal over that seen in Fig. 1). D primitive,  derived. 

Several interesting points emerge from these analyses. Many of the more-or-less 
controversial higher taxa (Cohorts and Orders) of Schram (1981), emerge, viz., Hemica- 
ridea (Cumacea, Tanaidacea, and Spelaeogriphacea), Brachycarida (Hemicaridea and 
Thermosbaenacea), Eucarida (Euphausiacea, Amphionidacea, and Decapoda), and 
Acaridea (Isopoda and Amphipoda). The latter also seems to bear some relationship 
to a yet unnamed new order being proposed by T. Bowman, R. Hessler, and H. Sanders 
which, interestingly, seems to fill one of the "unoccupied" 5aw/7/a>2£' of Schram (1981). 
On the other hand, some taxa derived from Schram (1981) do not seem viable: e.g., 


.& -^ 


Figure 3. Cladogram based on the same character data as that used in cladograms of Figure 1 , but analyzing 
only living groups and excluding the 4 extinct taxa, Palaeocaridacea, Belotelsonidea, Waterstonellidea, and 
Pygocephalomorpha. D primitive,  derived. 

Arthrostraca in the sense of Haeckel (1896), Giesbrecht (1913), or Grobben (1919) 
which unites all carapaceless syncarid and acaridean forms; or Mysoida (Belotelsonidea, 
Mysidacea, and Waterstonellidea), which seems invalid as a cladistic or taxonomic 

I decided to test the effect on the overall scheme of relationships when the fossil 
taxa were excluded from consideration (Fig. 3). Little change was noted except to ally 

/ / 

^ .V -^ .^^ $? 


fT7 i^ 


r J^ ^ ^ ^ I / J .^ ^5- j^ # ,o _^ / ^^ / ^^ ^s' 

^f .. 



/\/ //'// / / / // ^^ ^^^ / / // /\/ ^1 


Figure 4. Cladogram with all taxa and character data the same as in Fig. 1, except the scoring of character 
31 is reversed and the presence of a carapace is treated as a derived feature, as favored by Dahl (1983). D 
primitive,  derived. 

syncarids and eucarids as sister groups. Some slight modifications occurred in the 
arrangement of higher eucarids, but otherwise the basic relationships of the taxa of 
Fig. 1 are preserved. The total homoplasy value (a measure of the amount of con- 
vergence) is somewhat higher (388 vs. 372) in the non-fossil scheme than in that which 
includes the extinct groups, although the total length of both trees is not that much 
different, 44 without fossils as opposed to 46 with extinct groups included. 


I also tested the contention of Dahl (1983) that the carapace is a derived feature, 
the lack of a carapace being viewed as primitive (Fig. 4). This test resulted in a somewhat 
longer cladogram than those in Figure 1 (47 vs. 46), but one which has a dramatically 
higher total homoplasy value (510 vs. 372). Similar results were obtained when the 
data based on Dahl's concept were run without extinct taxa. It would seem, ♦therefore, 
that the suggestion by Dahl that the carapaceless state is the primitive one for eumalacos- 
tracans engenders a more complicated and less parsimonious array of relationships. 
Note, however, that the relationships within peracarids persist, including that of break- 
ing apart Mysidacea. 

Characters difficult to use 

Certain characters were deliberately not used here though they have found wide- 
spread employment in the taxonomy and phylogenetic discussions of Eumalacostraca 
by various authors. 

In eucarids, although the structure of maxillipedes was used (lamellate appendages 
with endites directed mediad), the number of them was not (three maxillipedes and 
thus the name decapod). The anatomical and functional state of thoracopods in higher 
eucarids is actually more varied than one would be led to believe from the automatic 
connotation engendered by the term "decapod." In several instances, e.g., many Den- 
drobranchiata, the so-called third maxillipede is actually more "pediform" in structure 
and function than "maxillipediform." Also, certain "pereiopods" actually have little 
locomotory function but are utilized in food acquisition and processing as well as 
defense. For example, in euzygids (=stenopodids) the fourth and fifth thoracopods are 
chelipedes and directed anteriad towards the mouth field resulting in a hexapodous 
condition instead of a decapodous one in these animals. In astacideans the characteristic 
great chelipedes of the fourth thoracopods serve in food procurement and defense, 
making the animal functionally octopodous. So while there are good maxillary and 
maxillipedal features which can serve to delineate a taxon Decapoda, ironically true 
decapody is not a particularly good character to assist in such delineation. 

Another feature taken for granted in discussion of eumalacostracan evolution is 
the fusion of the first thoracomere to the cephalon. Bathynellacea, of course, do not 
have this fusion. The Carboniferous genera Belotelson and Waterstonella apparently 
had free first thoracomeres as well, as they lacked maxillipedes altogether. Hence, it is 
imperative to resolve whether or not all living forms with a carapace do or do not fuse 
the first thoracomere to the head. For example, euphausiaceans lack a maxillipede, 
have the carapace fused to the thoracomeres, but have all thoracomeres associated 
together separate from the maxillary segment. Examination of mysidaceans revealed 
a variable state of affairs. Lophogastridans, with their well-developed maxillipedes, 
closely associate the first thoracomere with the cephalon and separate it from the second 
and following thoracomeres. However, in the mysidan Neomysis americana there is a 
separation of the maxillary from the thoracic segments, with all eight sets of thoracopods 
closely associated and separated by a distinct skeletal bar from the more anterior 
mouthparts. So in mysidans the first thoracomere is clearly not fused to the cephalon, 
although there is a tendency to develop maxillipedes. This feature serves to break apart 
the taxon Mysidacea, making Mysida a sister group to all other peracarids. 

I also excluded three characters which have been asserted as distinctly peracaridan, 
including the lacinia mobilis mentioned above. The presence of a manca stage is 
frequently cited as a characteristic of peracarids. Generally workers used this feature 
as if they were dealing with a manca larva. Mancas, however, are not to be equated 
with the zoea, cypris, or other larval types which have considerable cladistic merit (see 
for example character 2). A "manca" is a stage of development which can have various 
forms of expression (Newman 1983). Amphipods are generally said to lack a manca, 
yet some hyperiids are freed from the female in a virtual manca state (Laval 1980). 
Some adults express a permanent manca condition, e.g., the genus Thermosbaena. 


Manca stages also occur outside the peracarids, e.g., bathynellaceans hatch in an extreme 
"mancoid" condition lacking several of the posterior thoracopods and in the adults of 
some forms the last thoracopod can be missing or greatly atrophied. The presence or 
absence of a manca may be better understood in terms of constraints placed on de- 
velopment by egg size (e.g., Steele and Steele 1975). Characters of marsupial and 
maxillipedal form alone can be used to delineate peracarids more securely; and while 
the presence of a manca stage may assist in this delineation, it is not as unambiguous 
as one would suppose. 

The same observation can be made of the monocondylic coxa/basis articulation 
recently noted by Hessler (1982). This character might appear to be a useful congruent 
feature towards establishing a concept of Peracarida. However, it has a variety of 
expression difficult at this time to evaluate. For example, the monocondyle variously 
arises from positions that are either lateral (tanaids), purely posterior (Spelaeoghphus), 
or postero-lateral (all other peracarids). Nor do all thoracopods have this joint. In 
tanaids the third through fifth limbs have a dicondylic joint whereas only the sixth 
through eighth have the distinct monocondyle. Completely aberrant condyle, muscle, 
and/or joint arrangements are seen in amphipods and mysidaceans. These latter two 
groups also display different degrees of expression of these features throughout the 
whole thoracopodal limb series. Although coxal/basal structure seems to second per- 
acarid monophyly, problems with variety of expression and assessment of polarities 
between these variations preclude its use here. 

Characters rejected for use 

Two suites of traditional characters were completely rejected, viz., those of gut 
structure and embryo flexion which have played so prominent a role in the work of 
Siewing, and which resulted in the diametric separation of isopods and amphipods. 
The more that is discovered about gut morphology, the more it seems that the digestive 
system is too plastic to yield any useful data for phylogenetic analysis. Kunze (1981, 
and personal communication) has noted that the anatomy of the stomach of isopods 
is closely tied to feeding habits. Ide (1892) and Naylor (1955) provided details of gut 
structure in Idotea identical to that supposedly characteristic of amphipods, including 
an anteriorly directed mid-dorsal caecum in /. tricuspidata. Carol Diebel {pers. 
comm.) is finding that stomach structure among hyperiid amphipods is so diverse as 
to be uncharacterizable because of adaptations to particular feeding strategies. 

The other character rejected here, but given great weight by Siewing, is whether 
the developing embryo is flexed ventrally or dorsally within the egg membranes. First, 
few studies within and between groups of peracarids have been performed to determine 
the distribution of these states. Second, one of these flexures must be primitive and 
the other advanced. As such, only one of them can be used to characterize one of the 
groups which possesses it, but they are not both derived characters. It might appear 
that the dorsal flexure is derived, but insufficient data exist from within and without 
peracarids in order to assess polarity. And third, flexure in embryos seems better 
understood in terms of the mechanics of a particular developmental sequence rather 
than in terms of phyletic trends. Note that in forms with a ventral flexure, there is 
typically a very distinct egg-nauplius stage in early development, the development of 
the teloblasts lags behind that of the primary part of the head. The development of a 
caudal papilla and a caudal furrow which lead to ventral flexure is thus possibly related 
to the rapid development of the naupliar region. In contrast, in animals with a dorsal 
flexure, the appearance of the naupliar anlagen lags. In such forms the teloblasts not 
only appear early in the sequence of events around the blastopore, they initiate their 
divisions early such that the naupliar and anterior metanaupliar somites appear virtually 
simultaneously. It would appear that because of the slower head development the 
proliferation of body somites is allowed to occur along the entire ventral and posterior 
surfaces of the egg without the appearance of a caudal furrow or papilla to produce 
ventral flexure. Clearly the "phylogenetic power" of the apparent differences of flexure 
between isopods and amphipods has been somewhat overextended. 


Variant cladograms 

Two variants in the cladograms were produced by the program frequently enough 
to require some comment here. One is a variation in the higher decapods seen in Figures 
1 and 3. In one (Fig. 1), somewhat more parsimonious, eukyphids are placed as a sister 
group to euzygids and reptants. In the other (Fig. 3), reptants are a sister group of 
euzygids and eukyphids. The former is a more traditional arrangement, but the latter 
is all the more startling in light of the pregnant comment of Felgenhauer and Abele 
(1983) that it was their belief that the origins of the so-called "natant" groups of 
decapods ". . . are to be found among those groups traditionally considered reptants." 

Indeed the entire issue of relationships within the decapods is under intense study 
right now. Burkenroad (1981), using branchial and ontogenetic characters not employed 
in this analysis, essentially obtained an arrangement of taxa like that seen in Figure 3. 
However, Felgenhauer {personal communication) is examining various features of ex- 
ternal and internal anatomy of natant forms in an attempt to arrive at an assessment 
of cladistic relationships within decapods. For these reasons, it may be wise to avoid 
use of terms like Decapoda and/or Pleocyemata for the time being, and rather treat 
the taxa within Eucarida as one long transition series. 

Another notable variant is seen in the higher peracarids between Figs. 1 and 2. 
The scheme in Fig. 2 is slightly less parsimonious, but if one can tolerate the ad- 
ditional character reversal it entails, then the arrangement is a sequence of events which 
is of considerable biological interest. The isopod/amphipod line seems to represent one 
in which the thrust of the radiation is toward varied exploitation of food resources 
because of the great plasticity in gut structures. The brachycaridan line, especially when 
the pygocephalomorphs are associated with it, seems to be a line which represents 
exploitation of reproductive strategies. Both pygocephalomorphs and tanaids have 
cones on the males. The supposed seminal receptacles mentioned by Brooks (1962) on 
pygocephalomorphs bear little actual resemblance to such structures. These structures 
are more likely large genital cones on the eighth thoracic stemites of males. The 
brachycaridan line is generally characterized by respiratory specializations of the max- 
illipedes and thoracopods. It is also a transition series in which carapace, pleopods, 
and the oostegite brood pouch are reduced or lost, culminating in the condition seen 
in the thermosbaenaceans. Insofar as the component taxa are currently understood, 
this line also exploits reproductive and unusual sexual strategies that maximize the 
number of offspring from any one generation (e.g., see Sieg 1983, for tanaidaceans, or 
Corey 1981, for cumaceans). 


Several conclusions can be drawn concerning the analyses made here: 

1) At least in part, the taxonomic scheme for the Eumalacostraca suggested by the 
identification of Baupldne within the group (Schram 1981) is supported, especially in 
regards to peracarid types. The idea of a taxon Arthrostraca is not favored, but the 
reassociation of isopods with amphipods in the sense of the old taxon Edriophthalma, 
and the linking of short carapace forms, does have some merit. 

2) If some degree of uncertainty is accepted, then the relationships within Peracarida 
postulated by Slewing (1951, 1956) can be subscribed to, but only if considerable 
multiple convergences can be tolerated within a distinctly unparsimonious scheme. 

3) The concept of the carapace as a derived feature in the sense of Dahl (1983) is 
acceptable only by tolerating a great many more convergences than occur when the 
presence of a carapace is viewed as primitive. 

4) The concept of a formal taxon Mysidacea seems best abandoned. The three 
subtaxa traditionally placed within it (Lophogastrida, Mysida, and Pygocephalomor- 
pha) are distinct from each other regardless of whether the presence of a carapace is 
considered primitive or derived. 

What taxonomy of Eumalacostraca should be derived from all this? The eucarids 
are destined for some kind of realignment, especially of the higher taxa. The phylo- 


genetic integrity of the brachycaridans is stable enough, whether or not pygocephalo- 
morphs are closely associated with them. The resolution of relationships within the 
edriophthalman branch must await the description and evaluation of the new order of 
Bowman, Hessler, and Sanders, as well as a reevaluation of relationships within isopods 
and amphipods using a careful analysis of character states in all subgroups therein. In 
regards to the latter, we may resurrect the old taxon Laemodipoda, wherein caprellids 
and cyamids are separated as sister groups off by themselves. Such a study is currently 
under way. 

It is my intent here to point out two things. First, there is merit in carefully reflecting 
on what are the constituent structural plans expressed within any particular taxon, alert 
to the fact that any particular Bauplan may or may not be developed, or may or may 
not be the basis of an extensive radiation. Second, regardless of the ongoing philo- 
sophical and in large part tautological debate on taxonomic theory, we must make 
some organized careful evaluations of characters and what their condition and polarity 
might be throughout all members of a group. These are problems which have been all 
too often neglected in the history of crustacean studies, but are not unique to the study 
of these arthropods. 


Special thanks must be extended to Dr. Richard Brusca, San Diego Natural History 
Museum and Allan Hancock Foundation, for his valuable collaboration in the ongoing 
evaluation of character states in malacostracans. Without his expertise and encour- 
agement this study would not have been undertaken. Mr. Ernest Iverson, Allan Hancock 
Foundation, ran the various series of data sets with the Wagner 78 program using the 
computer facilities of the University of Southern California. Graphic work was done 
by Susan Mc Williams, San Diego Museum of Natural History. Valuable criticism of 
the manuscript has been offered by R. Brusca, G. Brusca, P. Delaney, M. Grygier, R. 
Hessler, and W. Newman. 

Literature Cited 

Brooks, H. K. 1962. The Paleozoic Eumalacos- 
traca of North America. Bulletins of American 
Paleontology 44:163-338. 

Burkenroad, M. D. 1981. The higher taxonomy 
and evolution of Decapoda. Transactions of 
the San Diego Society of Natural History 19: 

Caiman, W. T. 1904. On the classification of 
Crustacea Malacostraca. Annals and Magazine 
of Natural History (7)13:144-158. 

. 1909. Crustacea. In R. Lankester (ed.). A 

Treatise on Zoology. Vol. VII. Adam & Charles 
Black, London. 

Corey, S. 1981. Comparative fecundity and re- 
productive strategies in seventeen species of 
Cumacea. Marine Biology 62:65-72. 

Dahl, E. 1983. Malacostracan phylogeny and evo- 
lution. Crustacean Issues 1:189-212. 

, and R. R. Hessler. 1982. The crustacean 

lacina mobilis: a reconsideration of its origin, 
function, and phylogenetic implications. Zoo- 
logical Journal of the Linnean Society, London 

Felgenhauer, B. E., and L. G. Abele. 1983. Phy- 
logenetic relationships among shrimp-like 
decapods. Crustacean Issues 1:291-312. 

Fryer, G. 1964. Studies on the functional mor- 
phology and feeding mechanism of Monodella 
argentarii. Transactions of the Royal Society 
of Edinburgh 66:49-90. 

Giesbrecht, W. 1913. Crustacea. In Handbuch 
der Morphologic der Wierbellosen Tiere. Bd. 
4. Arthropoden. Jena: Gustav Fisher. 

Grobben, K. 1919. Uber die Musculatur des Vor- 
derkopfes der Stomatopoden und die syste- 
matische Stellung dieser Malakostraken- 
gruppe. Sitzungberichte der Akademie 
Wissenschaften, Wien 128:185-214. 

Haeckel, E. 1896. Systematische Phylogenie: 2 
der Wirbellose Thiere. Georg Reimer, Berlin. 

Hessler, R.R. 1969. Peracarida. Pp. R360-R363 
in R. C. Moore (ed.). Treatise on Invertebrate 
Paleontology, Part R., Arthropods 4, Vol. I. 
Geological Society of America and University 
of Kansas Press, Lawrence. 

. 1982. The structural morphology of walk- 
ing mechanisms in eumalacostracan crusta- 
ceans. Philosophical Transactions of the Royal 
Society, London (B)296:245-298. 

. 1983. A defense of the caridoid facies: 

wherein the early evolution of the Malacostra- 
ca is discussed. Crustacean Issues 1:145-164. 

Ide, M. 1892. Le tube digestion des Edrioph- 
thalmes; etude anatomique et histologique. 
Cellule 8:97-204, plates 1-7. 

Kunze, J. C. 1981. The foregut of malacostracan 
Crustacea: functional morphology and evolu- 
tionary trends. American Zoologist 21:968. 

Laval, P. 1980. Hyperiid amphipods as crusta- 
cean parasitoids associated with gelatinous 


zooplankton. Oceanography and Marine Bi- 
ology Annual Review 18:1 1-56. 

Naylor, E. 1955. The diet and feeding mechanism 
of Idotea. Journal of the Marine Biological As- 
sociation of the United Kingdom 34:347-355. 

Newman, W. A. 1983. Origin of the Maxillopoda: 
urmalacostracan ontogeny and progenesis. 
Crustacean Issues 1:105-120. 

Schram, F. R. 1974. Convergences between Late 
Paleozoic and modem caridoid Malacostraca. 
Systematic Zoology 23:323-332. 

. 1981. On the classification of Eumala- 

costraca. Journal of Crustacean Biology 1:1- 

1983. Method and madness in phylogeny. 

Crustacean Issues 1:331-350. 
Sieg, J. 1983. Evolution of Tanaidacea. Crusta- 
cean Issues 1:229-256. 

Slewing, R. 1951. Besteht ein engere Verwand- 
schaft zwischen Isopoden und Amphipoden. 
Zoologisches Anzieger 147:166-180. 

. 1956. Untersuchungen zur Morphologic 

der Malacostraca. Zoologisches Jahrbuch, Ab- 
teilung Anatomie 751:39-176. 

Steele, D. H., and V. J. Steele. 1975. *Egg size and 
duration of embryonic development in Crus- 
tacea. Internationale Revue der gesamten Hy- 
drobiologie 60:71 1-715. 

Watling, L. 1981. An alternative phylogeny of 
peracarid crustaceans. Journal of Crustacean 
Biology 1:201-210. 

. 1983. Peracaridan disunity and its bear- 
ing on eumalacostracan phylogeny with a re- 
definition of eumalacostracan superorders. 
Crustacean Issues 1:213-228. 



"^«»4p/^^ OF THE SAN DIEGO 

JUn society of 

1955 natural history 

Volume 20 Number 17 pp. 313-336 30 January 1985 

History and status of the avifauna of Isla Guadalupe, Mexico 

Joseph R. Jehl, Jr. 

Hubbs-Sea World Research Institute. 1700 South Shores Road, San Diego, California 92109, USA 

William T. Everett 

San Diego Natural History Museum, P.O. Box 1390, San Diego. California 92112, USA 

Abstract. Since 1954, renewed interest in Isla Guadalupe, stimulated by the research of the late 
C. L. Hubbs, has resulted in much new information on the avifauna. In this paper we review the 
status of the birdlife through 1982, provide an historical review of the research, including information 
on the timing and extent of the many expeditions, and provide a bibliography. 

Resumen. Desde 1 954, interes renovado en la Isla Guadalupe, promovido por las investigaciones 
del Dr. C. L. Hubbs, ha resultado en mucha nueva informacion sobre la avifauna. En esta presentacion 
detallamos lo conocido sobre la historia natural de los aves hasta 1982, damos un resumen historico 
sobre las investigaciones, incluyendo informacion sobre el estacionamiento y duracion de las expedi- 
ciones y ofrecemos una bibliografia. 


Because of its unique plant and animal life, Isla Guadalupe, Mexico, has fascinated 
biologists since the time of its scientific "discovery" in 1875. This rugged and remote 
volcanic island, 220 miles south of the Mexico-United States boundary and 160 miles 
west of the peninsula of Baja California, was the home of ten endemic species or 
subspecies of birds (an eleventh has recently been proposed). But the history of the 
birdlife "is a sad one of reduction and extermination through destruction of habitat by 
feral goats, predation by introduced house cats, and regrettably, some excess of zeal by 
collectors" (Howell and Cade 1954: see also Huey 1924, 1925), and many of the 
endemics are gone. 

Much has been written about the avifauna, and the history of some of the extinct 
forms has been well documented. Ridgway (1876), Bryant (1887(2), Thayer and Bangs 
(1908) and Hanna (1925) provided comprehensive reviews of the birdlife and these 
were made current by Howell and Cade in 1954. Since then, sufficient new information 
has been obtained to prompt a further compilation. 

For a description of the island and its general ecological settings and geology see 
Howell and Cade (1954), Lewis (1971), and Johnson (1953). A good summary is 
provided by Lindsay (1966:2); who wrote: "Guadalupe Island is about 22 miles long 
from north to south, and four to six miles wide over most of its length. The highest 
part is at the north end, where magnificent sheer cliffs tower over the sea or narrow 
beaches of cobbles and sand. The central part is a plateau sloping toward the south 
end, but the whole island is very rugged. Most of the base rock is red lava, and several 
of the lesser peaks are cinder cones. Recent research has shown that the oldest lava 
flows occurred about 7,000,000 years ago." 

Most of the island is devoid of vegetation. A remnant of a formerly large cypress 


forest (Cupressus giiadahipensis) about Vh miles long, is located in the north central 
part of the island. The endemic fan palm {Erythea edulis) is fairly common on the 
north slope of the island, and atop the main ridge at the north end there are stands of 
Island Oaks {Quercus tomentella) and Guadalupe Island Pines {Pinus radiata var. 
binata). The highest peak. Mount Augusta, rises more than 4200 feet abore the sea. 
A map of the island, with names of major localities, is given in Figure 1. 

History of Ornithological Research 

The Hungarian explorer Johan Xantus de Vesey is generally acknowledged as 
having been the first naturalist to visit the island. He was becalmed there on 1 7 March 
1859, while en route between San Francisco and Cabo San Lucas (Madden 1949). In 
his journals Xantus described several species of birds whose identity might be inferred. 
Yet, his accounts of the island itself are at such variance with those of an earlier French 
expedition (duPetit-Thouars 1956) that they undermine the credibility of his reports, 
and we have given them no attention. 

It was the work of the botanist Edward Palmer that drew first attention to the 
island. Palmer, collecting plant specimens for the U.S. National Museum, arrived in 
1875, intending to remain for six weeks (Blake 1961). Instead he found himself stranded 
for four months (Table 1), which afforded him ample time to collect eight of the nine 
endemic taxa of landbirds, all of which were quickly described by Robert Ridgway 

Palmer's visit came just in time, for the island was already undergoing a series of 
rapid and irreversible changes. Goats had been introduced as a source of meat by sealers 
or mariners, perhaps as early as the 18th century, and more were introduced for a 
commercial wool-producing enterprise in the ISlOs (San Diego Union, 15 March 1873; 
Anon. 1874). They numbered in the tens of thousands by the time of Palmer's visit, 
caused the elimination of much of the island's vegetation (already noted by the French 
in 1838), and forced much of the birdlife to be concentrated in the few wooded areas. 

Spurred by Palmer's discoveries, Walter E. Bryant spent a short time on the island 
in January 1885, then returned in December with the intention of spending six weeks 
there. Three and a half months elapsed before his ship returned. Palmer's work had 

Table 1. Chronology of ornithological research at Guadalupe Island, 1875-1953. 


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec 






Bryant 1885 


Anthony, Streator 


Gaylord, Anthony 


Anthony et al. 






Hartert, Rothschild ? 

Brown et al. 








Anthony et al. 




Hanna et al. 








Howell and Cade 1953 

Blake 1961 
Bryant 1887a 
Bryant 1887a 
Anthony 1901, 1925 

Gaylord 1897 

Davidson 1928, Kaeding 

Thobum 1899 
Abbott 1933 
Howell and Cade 1955 
Thayer and Bangs 1 908 
Townsend 1911, 1923 
Abbott 1933 
Kimball 1922 

Anthony 1925, Hanna 1925 
Huey 1924 
McLellan 1925 
Swarth 1933 
Huey 1954 

Bond and deSchauensee 1 944 
Howell and Cade 1954 






ROCK -o 




MT. AUGUSTA - 4257" 




29 00N 







00 — 


Figure 1. Isla Guadalupe, showing major localities and forested areas mentioned in this report. 

been so thorough that Bryant was unable to discover any additional endemic landbirds, 
but he did amass a number of unusual observations (Bryant 1 887a). More importantly, 
he discovered the Guadalupe Petrel {Oceanodroma macrodactyla), the first endemic 
seabird to be recognized (Bryant 1887/?). 

In the last decade of the 19th century, several expeditions made brief stops at 
Guadalupe. A. W. Anthony made several trips and was the first to report the destruction 
of biota by feral goats (Anthony 1901). Within a few years many of the endemic birds 
were extinct. The last report of a Guadalupe Caracara {Polyhorus plancus lutosus) was 
made by Rollo Beck, who collected nine in December 1900. An expedition from the 


Table 2. Expeditions to Guadalupe Island by C. L. Hubbs. 


Jan Feb 








Oct Nov Dec 

Total days 






* 8 














































Thayer Museum, including W. W. Brown, spent two months on the island in the spring 
of 1906. After scouring the island, they declared the Guadalupe Wren {Thryomanes 
bewickii brevicauda) and Guadalupe Towhee (Pipilo erythrophthalmus consobhnus) 
extinct (Thayer and Bangs 1908), and predicted the quick demise of the Guadalupe 
nicker (Colaptes auratus rufipileus). Yet, despite intense predation by cats, the Gua- 
dalupe Petrel persisted "in large numbers." Only six years later, however, in 1912, 
Beck collected two downy chicks, the last known examples (Davidson 1928). 

After the basic collections had been made, and the extinction of the endemics had 
been documented, interest in the island lagged. Some ornithologists continued to visit 
it (often en route to more exciting localities) in the 1920s and 1930s (e.g., Hanna 1925), 
but little new information resulted. 

Commencing in the 1940s and continuing through the early 1970s studies by Carl 
L. Hubbs and his many colleagues renewed interest in Guadalupe. Between 1946 and 
1971, Hubbs made 21 expeditions to the island (Table 2). Hubbs discovered that small 
islets at the southern end of the main island were important seabird colonies. In addition 
to initiating population studies on seabirds, he reported the apparently distinctive 
summer and winter populations of Leach's Storm-Petrels (Hubbs 1960) that have 
received much subsequent attention (Crossin 1974, Ainley 1980, 1983, Bourne and 
Jehl 1982). He also discovered the only known fossil locality on the island (Pleistocene), 
which has yielded the remains of a few seabirds (Hubbs and Jehl 1976). 

In 1953, T. R. Howell and T. J. Cade spent five days on the island, on one of 
Hubbs' expeditions. Their findings (Howell and Cade 1954, 1955) resulted in the first 
discussion of the birdlife in several decades. Additional work by the Pacific Ocean 
Biological Survey Program (POBSP) in the late 1960s and early 1970s (DeLong and 
Crossin MS, Brownell MS, Crossin MS, 1 974) revealed much about the nesting seabirds. 

Since the late 1960s we have each had occasion to visit Guadalupe several times. 
Our observations, and those of many other biologists who have generously contributed 
data (Table 3), and the voluminous field notes of the late Carl L. Hubbs provide the 
basis for this report. Note, however, that as with earlier studies nearly all of the recent 
work has been concentrated in late winter through early summer and that the fall 
season, with one exception, is unrepresented. We have attempted to make the report 
comprehensive and current, and to provide a complete ornithological bibliography. 
This compilation was stimulated in part by a request, for planning purposes, from the 
Mexican government, through Dr. Martin Gonzales. Because prospects of additional 
surveys in 1982 and 1983 failed to materialize, we are making the report available 
now. We acknowledge, however, that much remains to be learned before our knowledge 
of the present avifauna is adequate. 


Table 3. Recent ornithological observations on Isla Guadalupe, Mexico. 





27-31 Jan 


26. 27, 30 Apr, 
1^ May, 
21-24 Oct 


17-22 Apr, 
20-23 Jun, 
28-29 Jun 


21-23 Feb 


19-27 Jan, 
12-18 Apr 


1 5 Mar, 
17-23 May 


mid Dec 


22 Nov-26 Dec 


12-13 Jul 


20 Jun-13 Jul 


9-20 Feb 
16 Apr 
17-18 Dec 


24 Mar, 18 Apr 


4-5 May 


1 9-26 Aug 


12-15 Feb, 
4-6 Mar 

K. L. Kenyon 

R. L. DeLong and R. S. Crossin 

R. L. Brownell, R. S. Crossin, R. L. DeLong 

J. R. Jehl, Jr. 
J. R. Jehl Jr. 

J. R. Jehl, Jr. 

K. Briggs 

E. Mirsky 

J. R. Jehl, Jr. 

M. Pierson and M. Riedman 

M. Pierson, E. Chu et al. 
W. T. Everett 
K. Briggs 

W. T. Everett 

W. T. Everett 

D. Duncan 

R. Condit 

Annotated List of Species 

In the following section we provide synoptic information on all species of birds 
recorded on Guadalupe Island and in waters immediately adjacent (within approxi- 
mately 1 5 km) of the island. We include information on many species of seabirds, 
which were not considered by Howell and Cade (1954). Nomenclature follows the AOU 
Check-list (1983), except for Pluvialis fulva (see Connors 1983). The following abbre- 
viations are used: AMNH, American Museum of Natural History; CAS, California 
Academy of Sciences; HSWRI, Hubbs-Sea World Research Institute; LACMNH, Los 
Angeles County Museum of Natural History; SDNHM, San Diego Natural History 
Museum; UCLA, University of California Los Angeles. 

Arctic Loon (Gavia arcticd) 

Uncommon winter visitor. Palmer found an adult washed ashore on 20 May 1875 
(Ridgway 1876). Hubbs collected one on 13 February 1957 and saw two others on his 
expedition (8-15 February 1957), and Mirsky (MS) observed one on 23 November 

Pied-billed Grebe {Podilymhus podiceps) 

Accidental. Hubbs collected one on 27 October 1957; location of specimen un- 

Eared Grebe {Podiceps nigricollis) 

Considered accidental by Howell and Cade (1955) on the basis of a specimen 
reported by Rothschild and Hartert (1902). There are four subsequent reports. Hubbs 


Figure 2. Guadalupe Storm-Petrel {Oceanodroma macrodactyla). Photograph of a mount in the Field 
Museum of Natural History, Chicago. 

collected two on 12 February 1957 (LACMNH) and reported taking another on 30 
October 1957; Mirsky (MS) reported one on 23-25 November 1973. 

Western Grebe (Aechmophorus occidentalis) 

Accidental. Two reports by Hubbs of single birds on 12 November 1954 and 17 
December 1957. 

Short-tailed Albatross {Diomedea albatrus) 

Probably regular in the area until the late 19th century. Bryant (1889) reported 
five between Ensenada and Guadalupe in April 1886. Now very rare and not reported 
in Baja California for many decades. 

Black-footed Albatross {Diomedea nighpes) 

Uncommon but regular near the island through most of the year, with many sight 
records from September to June. 

Laysan Albatross {Diomedea immutabilis) 

This species is a rare but regular wanderer to Baja California. R. Wisner observed 
a single bird about 3 km off the north end of the island on 22 April 1958, and R. 
Pitman saw one 90 km to the northeast on 4 January 1980. Anthony (1898Z)) took a 
specimen between San Geronimo and Guadalupe islands in March 1897. 


Northern Fulmar {Fulmams glacialis) 

Irregular visitor in winter. Mirsky (MS) reported one on 6-7 December 1973. This 
species reaches the latitude of central Baja California in flight years. 

Cook's Petrel {Pterodroma cookii) 

This species occurs regularly off' the coast of Baja California (Jehl, pers. obs.). Two 
were seen on 1 1 April 1961, 100 km north of the island (Small 1961). Pitman saw one 
along with another unidentified Pterodroma within 15 km of Guadalupe on 1 1 October 

Pink-footed Shearwater (Puffinus creatopus) 

According to Grinnell (1928): "First definitely recorded by Gaylord ... as seen by 
A. W. Anthony near Guadalupe Island, September 17, 1896." While this record may 
be correct— the species is a common migrant nearer shore— we suspect that it more 
likely pertains to the Black- vented Shearwater, which breeds locally. 

Sooty Shearwater {Puffinus griseus) 

Probably a regular visitor from April to October. Thayer and Bangs ( 1 908) reported 
taking two specimens "near" the island in June 1906, and Pitman saw one on 4 January 

Black-vented Shearwater (Puffinus opisthomelas) 

This shearwater has long been known to breed at Guadalupe. Anthony (1900) 
reported it to be "rather common ... in several parts of the island, but in no place 
was there any large colony. Their nests were all inaccessible owing to the nature of the 
sites selected, whether in natural holes in the lava or under large boulders, and no eggs 
were secured. A night was spent on the top of the island in a heavy cypress growth, 
about 4000 feet above the sea. Here the shearwaters were heard all night, their choking, 
gasping notes coming from all sides as they flew through this grove." 

Hubbs discovered and for many years surveyed the colony on Islote Negro, which 
in the late 1960s and early 1970s contained 100-150 pairs (Jehl, pers. obs., Crossin 
MS, Brownell MS). Crossin visited Islote Afuera on 22-23 June 1968 and found a 
colony of 150+ pairs. We suspect that there may be other colonies on the main island 
because in the early evening flocks of 500-2500 shearwaters, too many to be accom- 
modated by known nesting areas, stage at the south end of the island. 

The breeding season at Islote Negro can be outlined with some confidence. By late 
November single birds begin to occupy burrows. By early January pairs are commonly 
found and fresh eggs are present by 5 March (Hubbs field notes). The peak of the egg- 
laying season occurs in early April, when most burrows contain an egg, but may extend 
to late June (Jehl, pers. obs.; Crossin MS). Young are present as early as late April and 
in late June most burrows have a chick. There are no data for later in the fall. Crossin 
reported that the phenology of the Islote Afuera colony was somewhat later than at 
Islote Negro. 

Leach's Storm-Petrel {Oceanodroma leucorhod) 

There has been much uncertainty about the historical status of storm-petrels at 
Guadalupe, much of which may never be fully resolved because the early literature is 
inconsistent, and because cats have eliminated most, if not all, storm-petrel colonies 
on the main island, whose location and species composition cannot be fully recon- 
structed. The complicated case of the Leach's Storm-Petrel is best treated chronolog- 

In the winter and spring of 1885-86, W. E. Bryant was stranded on Guadalupe 
for nearly four months. Near the end of his stay he discovered storm-petrels nesting 


among the pines and cypress trees at the northern end of the island. Bryant (1887a) 
reported these as Leach's Storm-Petrels {Oceanodroma leucorhoa), and recognized that 
they had a more deeply forked tail and a longer middle toe than other races. Bryant 
(1887^) then named these as a new race {O. I. macrodactyla), the Guadalupe Storm- 
Petrel, which the AOU (1889) elevated to species rank. 

In 1889, C. H. Townsend secured a single small storm-petrel, which had the sides 
of the rump "whitish," at sea near Socorro Island, 1200 km SE of Guadalupe. He 
described it as a new species {O. socorwensis), guessing that it nested at Socorro Island 
(Townsend 1890). 

A. W. Anthony visited Guadalupe in May 1892 and collected three nestling Gua- 
dalupe Storm-Petrels, but did not mention other species. In 1896, Anthony and H. A. 
Gaylord visited Guadalupe in mid-September. Anthony spent a night atop the island 
but reported no petrels of any species (Anthony 1898c). However, Gaylord (1897) 
stated: "Regarding the Petrels which breed on the island, the hunters told us that 
while doing some stone work in the region of the Petrel colony, they had found two 
different species. They described the Guadalupe Petrel and an entirely black one, which 
together with a wing found on the trail to the cypress grove makes it appear that O. 
homochroa is an inhabitant of the island." 

In late March, Anthony collected a series of adult macrodactyla, along with a few 
eggs. That summer he returned and collected young, noting that macrodactyla leaves 
the colony by 10 June. On neither trip did he find other species in the colony, but in 
June he collected a storm-petrel with enlarged ova at sea 120 mi north of Guadalupe, 
which became the type of Kaeding's Petrel {Oceanodroma kaedingi). This new form 
was distinguished by its small size and white rump, but Anthony (1898a) noted that 
there was much variation in rump color. From observations at sea, Anthony (1898a) 
surmised that the range of kaedingi extended from the Revillagigedos to southern 
California and guessed (1898c) that its breeding grounds were on Guadalupe. Also, he 
reidentified the wing ascribed to homochroa in 1896 as belonging to kaedingi (Anthony 

H. B. Kaeding (1905), who accompanied the 1897 expedition, was the first to allege 
a common breeding area for kaedingi and macrodactyla. He wrote that "the breeding 
grounds of kaedingi are as yet unknown, but it is probable that the birds occupy the 
burrows of Guadalupe Petrels . . . after the breeding season of the former is closed." 
In late May 1906, W. W. Brown and H. W. Marsden visited the breeding grounds of 
the Guadalupe Storm-Petrel (Thayer and Bangs 1908); they found O. macrodactyla 
but no other species. 

By this time most of the islands off Baja California had been surveyed and tax- 
onomists began to re-evaluate the variation in the Leach's Storm-Petrel group (including 
Swinhoe's Storm-Petrel, O. monorhis, of the western Pacific), a process that still con- 
tinues (e.g., von Berlepsch 1906, Emerson 1906, Godman 1907-1910, Obserholser 
1917,Loomis 1918, van Rossem 1942, Austin 1952, Todd 1955, Palmer 1962, Crossin 
1974, Ainley 1980, 1983, Bourne and Jehl 1982, Power and Ainley MS). The work 
was made difficult because of the petrels' discontinuous breeding range, the scarcity of 
specimens from breeding colonies, the great variability in some populations, and the 
lack of knowledge about the location of breeding grounds. The last point was illustrated 
by A. C. Bent (1922), who reported that "Mr. Anthony wrote me that on Guadalupe 
Island the Guadalupe Petrels breed early, April 20 or earlier, and that after they are 
through breeding the Kaeding Petrels use the same burrows." As Anthony was the most 
knowledgeable ornithologist regarding the petrels of the region no one questioned his 
perpetuation of Kaeding's (1905) idea, even though all expeditions to the nesting grounds 
of macrodactyla failed to report any other species as nesting, and despite the fact that 
the nesting grounds of kaedingi had yet to be discovered (e.g., Oberholser 1917, Grinnell 

On 11-17 July 1 922 Anthony (1925) returned to Guadalupe and visited the nesting 
area of macrodactyla. That species had disappeared, but the expedition did find nu- 
merous bodies of kaedingi that had been killed by cats or were impaled on cholla, and 


even found a week-old chick (CAS no. 25561). Unfortunately, no data were published 
on precise localities. Hanna (1925) stated that kaedingi "lived among the loose rocks 
and in holes in the cliffs," whereas Anthony (1925) said that it "evidently" nested in 
high cliffs at the north end of the island. Regardless, the discovery of the chick (i) 
provided the first proof that a species other than macrodactyla nested on the main 
island, (ii) established the island as a breeding location for kaedingi, (iii) is in accord 
with recent data that kaedingi is a crevice nester {macrodactyla nested in burrows in 
forested areas), and (iv) shows that Anthony had no evidence of burrow-sharing between 
macrodactyla and kaedingi. 

In 1950, Carl L. Hubbs discovered and began to study the Leach's Storm-Petrels 
nesting on small islets at the south end of the main island. Hubbs (1960) was the first 
to recognize that there were evidently two populations, a larger form with white rump 
that bred in winter and fiedged young by April, and a smaller form with less white on 
the rump that began laying in June and fledged young in October or November. Surveys 
by the POBSP in June 1968 determined the summer population on Islote Negro at 
4000 adults and on Islote Afuera at 3000 adults (Crossin MS, 1974). Jehl also found 
a nearly-grown chick on Gargoyle Rock in April 1970 and Huey (1952) reported a 
chick there in January 1950. Huey also reported that Hendrickson collected "a number 
of Oceanodroma petrels from rock crevices on the hillsides of Melpomene Cove, sit- 
uated on the southern end of Guadalupe Island." We presume that small colonies 
remain to be discovered on the main island but none are known with certainty. Evidence 
of the persistence of one or more colonies there is provided by repeated observations 
of birds landing on ships at the north end of Guadalupe in late spring and summer 
(Jehl, pers. obs., Anthony 1925, Huey 1930), as well as records of a few petrels flying 
over the north end of the island in May 1971 (Jehl 1972) and many near the cypress 
grove in August 1981 (R. Moran, D. Duncan, pers. comm.). 

The correct nomenclature of the Guadalupe populations has been debated. Van 
Rossem ( 1 942) pointed out that the type of socorroensis was a small bird of the Gua- 
dalupe population; since that name had priority, kaedingi was relegated to synonymy. 
When Hubbs (1960) found that two populations were present on Islote Negro, he 
referred to the summer breeders as socorroensis and the winter breeders as kaedingi. 
However, the situation is evidently even more complicated, because the summer birds 
are highly variable. Crossin (1974) reported that all birds nesting at Islote Negro were 
dark-rumped, while on Islote Afuera over 90% of the population was white-rumped. 
Recently, Ainley (1980) named the winter breeders as a new race, cheimomnestes, on 
grounds unconvincing to Bourne and Jehl (1982) but reaffirmed by Power and Ainlev 

In summary. Leach's Storm-Petrels breed commonly on islets at the southern end 
of Guadalupe Island. There is also strong presumptive evidence for the persistence or 
re-establishment of colonies near the north end and center of the main island. According 
to Power and Ainley (MS), the winter and summer populations on Islote Negro are 
temporally segregated and morphologically distinct. If so, their findings would require 
that the two populations be considered as distinct species, not subspecies. Yet, because 
of the lack of field work in the fall, neither Bourne and Jehl (1982) nor Crossin (1974: 
1 74) were convinced that the populations are fully isolated temporally. Furthermore, 
because there are differences in coloration (at least) between summer populations breed- 
ing contemporaneously on I. Negro and 1. Afuera. and because the morphological 
characters and breeding seasons are not known for the presumed colonies on the main 
island, the situation remains unclear. This is one of the most complicated cases of 
differentiation known in birds, and one that will require additional study. 

Ashy Storm-Petrel {Oceanodroma homochroa) 

A wing attributed to this species by Gaylord (1897) was reidentified as that of O. 
leucorhoa by Anthony (1898r). In Gaylord's original report (see above) the presence 
of an "entirely black" storm-petrel near the colony of O. macrodactyla on the main 


island was alleged by "hunters." Probably these represented dark-rumped examples of 
the summer population of 6>. /. socorroensis or, less likely, wanderers of O. /. chapmani 
from the San Benito Islands. O. homochroa may occur off Baja California in winter, 
but the nearest breeding colony is on Los Coronados, 320 km to the north, and may 
consist of only 2-3 pairs (Jehl pers. obs.). , 

Wedge-rumped Storm-Petrel {Oceanodroma tethys) 

A specimen obtained in 1950 was said to have been collected from a crevice on 
the side of Melpomene Cove, at the south end of the main island, which also included 
a downy young of 6>. leucorhoa (Huey 1952). However, the precise locality is not clear. 
Howell and Cade (1954) claimed it was collected on Gargoyle Rock at the end of the 
island, and Hubbs told Jehl the same thing. Jehl landed there in April 1970 and found 
a single chick of O. leucorhoa. There are no subsequent records although the species 
occurs regularly to southern Baja California (Jehl pers. obs.. Pitman pers. comm.). 

Black Storm-Petrel {Oceanodroma melania) 

Kaeding (1905) reported this species near Guadalupe, but the observation could 
have pertained to any of several others. Anthony's (1898c) report, which implies that 
the species nests on Guadalupe Island, actually alludes to the nesting season of the 
species elsewhere. 

Guadalupe Storm-Petrel {Oceanodroma macrodactyla) 

This endemic and unusual storm-petrel (Fig. 2), originally described as a race of 
O. leucorhoa (Bryant 1887^), bred in soil burrows atop the main island, among the 
pines at the north end and in the cypress grove. Its status and history have most recently 
been reviewed by Jehl (1972). Both Howell and Cade (1954) and Greenway (1967) 
incorrectly cite the last report of the species, apparently overlooking the work of Da- 
vidson (1928), who corrected earlier errors and established August 1912 as the last 
acceptable record. A winter breeder, it laid by early March (Kaeding 1905) but some- 
times as late as June, for Beck collected three chicks on 3 August 1912 (specimens in 
AMNH). Nothing is known about the ecology or distribution of this species at sea. 
Thobum (1899) reported it as abundant about his ship at night in late June 1897, as 
it lay anchored at Guadalupe, and reported collecting several. If so, these would rep- 
resent the only documented records of macrodactyla away from the breeding colonies, 
but we have been unable to locate any specimens and suspect that his reports refer to 
O. leucorhoa. 

The Guadalupe Storm-Petrel was considered abundant in the colony as late as 
1906 (Thayer and Bangs 1908), but was being preyed upon heavily by domestic cats. 
Davidson (1928), in declaring it to be extinct, based her conclusion on the negative 
results of the 1922 expedition by the Cahfomia Academy of Sciences (Hanna 1925), 
which took place in mid-summer, after the main breeding season, and on the erroneous 
assumption that the CAS expedition of April 1925 (McLellan 1926) had carefully 
searched the breeding grounds. Jehl (1972) spent several nights listening for petrels in 
the early 1970's in the pine-oak woodlands atop the island, but did not visit the cypress 
grove, where the species also had nested. No thorough survey of the breeding grounds 
has been made at the appropriate season since 1906. The apparent persistence of <9. 
leucorhoa on the main island despite predation by cats allows some hope that macro- 
dactyla may still exist. 

As noted above there is no acceptable evidence for the often-repeated contention 
that Guadalupe and Leach's storm-petrels ever bred in the same colonies. All evidence 
suggests that they used different habitats: macrodactyla burrowed in soil, leucorhoa 
nested in crevices. Further, the breeding season of macrodactyla probably overlapped 
that of the winter and summer forms of leucorhoa, so that sequential use of burrows 
would have been impossible. 


Red-billed Tropicbird {Phaethon aethereus) 

Probably regular in the area but the only records are of individual birds 25 and 
37 km north of the island on 1 1 October 1979, and another slightly farther north on 
7 January 1980 (Pitman). 

Red-tailed Tropicbird {Phaethon rubricauda) 

Anthony (1898Z^) reported collecting a specimen close to Guadalupe on 23 July 

Brown Pelican {Pelecanus occidentalis) 

There are two records, an immature individual reported by Anthony (1925), and 
a probable juvenile reported on 4 July 1977 (Pierson and Riedman, MS). This coastal 
species rarely wanders to deep waters beyond the continental shelf. 

Double-crested Cormorant {Phalacwcorax auritus) 

Probably a rare or accidental visitor, but its status requires verification. One was 
reported by Gaylord (1897) and Huey (1924) casually mentions the species as being 
present in 1923. Hubbs reported the species on several trips and on 11 June 1955 
described a cormorant with "bill and pouch yellow" that would seem to be this species. 

Brandt's Cormorant (Phalacwcorax penicillatus) 

Resident. Seen regularly in small numbers along the entire east side of the island 
but commonest near the southern end, where a few pairs breed on outer islet (McLellan 
1926, Crossin MS), and on Islote Zapato (Hubbs notes, Jehl pers. obs.). The maximum 
single count is 20 at I. Zapato on 4 May 1966. We suspect that the entire island 
population does not exceed 30-40 individuals. Specimens in SDNHM. 

Pelagic Cormorant {Phalacwcorax pelagicus) 

J. Sefton reported this species on Hubbs' expedition of 27 January-3 February 
1950 (Hubbs field notes). His identifications were doubted by Hubbs, and by us. 

Magnificent Frigatebird {Fregata magnificens) 

Sightings of an immature on 1 and 4 July 1973 (Pierson and Riedman, MS) 
probably represent the same individual. 

Great Blue Heron {Ardea herodias) 

Probably a rare but regular winter visitor. Hubbs saw one or more on five different 
trips (maximum three, two trips), between November and February. Other records are: 
1, September 1896 (Gaylord 1897); 2-3 in summer 1922 (Anthony 1925); and 1, 
midwinter 1965 (Kenyon MS). 

White-fronted Goose {A nser albifrons) 

Bryant (1887(3) shot one on 14 January 1885, but it fell over a cliff' and could not 
be recovered. 

Brant {Branta nigricans) 

Mirsky (MS) reported a sick bird at Northeast Anchorage on 22-24 November 


Mallard {Anas platyrhynchos) 

Hubbs saw several and collected single males on 13 and 17 December 1957 

Northern Pintail {Anas acuta) 

Pitman saw one 40 km north of the island on 1 1 October 1979. 

Blue- winged Teal {Anas discors) 

Hubbs reported collecting an adult male on 30 October 1957. The location of the 
specimen, if preserved, is unknown. 

Cinnamon Teal {Anas cyanoptera) 

Jehl saw a male swimming along the shore of the main island on 21 January 1970. 

Lesser Scaup {Aythya affinis) 

Hubbs' field notes list "a female or immature male" several kilometers from the 
island on 22 November 1964. 

Red-breasted Merganser {Mergus senator) 

This species probably is an occasional winter visitor. There are two records: 28 
January 1950 (Hubbs), and 13-17 December 1973 (Mirsky MS). 

Osprey {Pandion haliaetus) 

Status uncertain. Ospreys have nested on many islands along the Baja California 
peninsula and perhaps formerly bred on Guadalupe, although proof is lacking. Speci- 
mens were collected 1 1 July 1922 (Anthony 1925, Hanna 1925) and on 25 July 1941 
(Bond and Meyer de Schauensee 1944). Kenyon (MS) visited Guadalupe early in 1965 
and saw no Ospreys but reported two presumed nests near the north end of the island. 
There are no other reports or indications of the species' presence. 

Red-tailed Hawk {Buteo jamaicensis) 

Formerly resident in small numbers. Howell and Cade (1954) considered it "ap- 
parently resident until at least 1932," but none of the early explorers were able to find 
any nests. Palmer {in Bryant 1887a) considered it as common as the caracara. Thayer 
and Bangs (1908) and Anthony (1925) reported that three or four could be seen in a 
day; Hanna (1925) also considered it common. This hawk wanders to many offshore 
islands in fall migration and probably reaches Guadalupe infrequently. However, we 
know of no recent reports for any season, and it is not resident at this time. 

Crested Caracara {Polyborus plancus lutosus) 

Extinct; formerly resident in small numbers. The detailed history of this endemic 
form and its taxonomy have been reviewed by Abbott (1933) and Brown and Amadon 

American Kestrel {Falco sparverius) 

Resident in small numbers. Bryant (1887a) stated that they were found most often 
in the central and higher portions of the islands. Howell and Cade (1954) reported a 
pair with young on a cliff' overlooking the sea at Northeast Anchorage; other birds were 
in the area. In recent years the species has been seen regularly near Northeast Anchorage 
and at the southern end of the island. D. A. Duncan {pers. comm.) visited Guadalupe 
19-26 August 1981 and reported it as common everywhere, one or two being seen at 


most localities. The Guadalupe population was described as an endemic race (gua- 
daliipensis) by Bond (1943); its validity was accepted by the AOU Check-list (1957) 
though not by the Mexican Check-list (Friedmann, Griscom, and Moore 1950). 

Peregrine Falcon {Falco peregrinus) 

This large, maritime falcon is likely to have occurred regularly during migration, 
but we know of only one report, a single bird seen on 19 September 1896 (Gaylord 

Prairie Falcon {Falco mexicanus) 

Br>'ant (1889) reported that the species was seen on "two or three occasions" in 
1886, but we suspect that these sightings pertain to the Peregrine Falcon. 

Pacific Golden Plover (Pluvialis fulva) 

Jehl saw a flock of 20, two km north of the settlement at the south end on 22 
February 1969. Mirsky (MS) reported from 1 to 12 birds along the shore at Northeast 
Anchorage from 23 November-16 December 1973. Presumably all records of golden 
plovers pertain to this species {see Connors 1983). 

Killdeer {Charadrius vociferus) 

Hubbs reported two on 13 December 1957. 

Willet {Catoptrophorus semipalmatiis) 

The only report is a single bird observed between 10-14 February 1977 (E. Chu). 

Wandering Tattler {Heteroscelus incanus) 

This is a regular visitor to the island from fall through spring; there is one summer 
record. One or two, often more, are seen on most trips. 

Ruddy Turnstone {Arenaria interpres) 

Though not reported by Howell and Cade (1954), this species is a regular visitor 
in small numbers. There are specific records for June, November-January, and April. 
At least three were present in November 1964 (Hubbs). Hubbs also collected several 
specimens, the location of which is not known. 

Black Turnstone {Arenaria melanocephala) 

Uncommon but regular in migration and during the winter. There are records for 
October-February, and April. The maximum count is seven on 20-26 January 1970 
(Hubbs, Jehl). Specimen LACMNH. 

Sanderling {Calidris alba) 

Two were seen on 22 January 1970 (Jehl). 

Western Sandpiper {Calidris mauri) 

One was photographed at Northeast Anchorage on 16 April 1978 (Everett). 

Short-billed Dowitcher {Limnodromus griseus) 

Hubbs collected an immature that landed on his boat about 2 km off the south 
end of the island on 29 August 1956 (LACMNH). 


Common Snipe {Gallinago gallinagd) 

One record, atop the northern end of the island on 8 June 1953 (Howell and Cade 

Red Phalarope {Phalaropus fulicaria) 

Regular in migration. This phalarope is seen irregularly, sometimes in fair numbers, 
between November and May; it has also been reported in late June (Thayer and Bangs 

Jaegers {Stercorarius spp.) 

Jaegers certainly occur near the island during migration, but the only published 
record seems to be that of Gaylord (1897), who reported two Long-tailed Jaegers {S. 
longicaudus) on 17 September 1896. Pitman has seen several jaegers in the area in 
January, and identified a Pomarine {S. pomahnus) on 4 January 1980. 

Heermann's Gull {Larus heermanni) 

Two adults were photographed by S. Leatherwood in January 1 973 (photo HSWRI). 

Ring-billed Gull {Larus delawarensis) 

Jehl and R. DeLong saw one immature at Northeast Anchorage on 22 January 
1970. This species rarely ventures beyond the coastal beaches. Hubbs reported "a few" 
on 28 January 1950, but his identification seems questionable. 

California Gull {Larus californicus) 

Though not recorded by Howell and Cade (1954), this gull is a regular, sometimes 
common, winter visitor. It avoids the elephant seal beaches, because of competition 
with the larger gulls, and tends to occur at sea. Twenty in February 1978 (Chu et al. 
MS) is the largest number recorded (but see Herring Gull). 

Herring Gull {Larus argentatus) 

Common winter visitor from November-April, at times being as common as the 
Western Gull. Hubbs reported that it was by far the commonest gull in January- 
February 1950, and counted 360 at the south end of the island in late January 1960. 
However, we suspect that many of these were California Gulls, for at that season Herring 
Gulls congregate near the elephant seal rookeries. 

Thayer's Gull {Larus thayeri) 

Uncommon but regular winter visitor. There are several records, all for immature 
or sub-adult birds; 21 February 1969, 16 April 1970, 30 January 1971 (3) and 15 
March 1971 (Devillers et al. 1971). 

Western Gull {Larus occidentalis) 

This species is resident at Guadalupe. Hubbs recognized that the local population 
differed slightly from the mainland birds. There are minor differences in the color of 
the fleshy parts (Howell and Cade 1 954) and also in the pattern of the primary markings. 
Hubbs (1960) suggested that it might represent an endemic race, but no formal analysis 
of the variation has been attempted. In winter the local population is probably enhanced 
by representatives from the mainland, as both Hubbs and Jehl have seen many birds 
with pinkish (rather than whitish) legs at that season. 

In Jehl's opinion the population in 1969-71 consisted of only 30-40 pairs. Crossin 
(MS) reported that the species is "rather sparse," and guessed that the local population 


in June was no larger than 200 birds. Jehl found a nest with three eggs near the old 
Lobster Camp on 21 May 1971, and Crossin reported another on Islote Afuera on 20- 
23 June 1968. In contrast to mainland gulls, the Guadalupe birds nest singly, well back 
from the shore, and there is no evidence of colonies. In November-December 1973, 
Mirsky reported 100 at the Northeast Anchorage; all had whitish legs. Chu et al. (MS) 
counted 100-125 along the entire eastern shore of the island in February 1978, and 
noted that adults outnumbered juveniles by about 10:1. Pierson and Riedman (MS) 
reported at least 100 birds during a circumnavigation in the first week of July 1977, 
most of which were attending nests; at least 15 large nestlings were seen. 

Bryant (1887a) was told that gulls nested commonly at the southern end of the 
island, ''where they were not so frequently molested by the 'Quelelis'" (=Caracaras). 

Glaucous-winged Gull {Larus glaucescens) 

Regular winter visitor, most frequently reported at the Northeast Anchorage in 
January-March, when they and other gulls feed on elephant seal remains and placentas. 
Up to 25, adults and immatures, have been seen at that time (Kenyon, MS). There are 
records from November-May, the latest being 1 May 1967 (Hubbs). 

Black-legged Kittiwake (Rissa tridactyla) 

The Kittiwake occurs in winter; it is common in some years, absent in others. 
Flocks of up to 100 were seen around the island on 20-26 January 1970 by Hubbs and 

Sabine's Gull {Xema sabini) 

This migrant is probably uncommon but regular in spring and fall. Hubbs reported 
10 birds 5 km east of the island on 26 April 1967. An additional report, on 27 January 
1950 (Hubbs) almost certainly pertains to an immature kittiwake. 

Royal Tern {Sterna maxima) 

Gaylord (1897) reported one near the island on 17 September 1886. 

Arctic Tern {Sterna paradisaea) 

This species certainly occurs regularly off the coast of Baja California, but there 
are few records. Pitman identified one near the island on 1 1 October 1979 and saw a 
second tern, probably of the same species. 

Xantus Murrelet {Synthliboramphus hypoleuca) 

This small alcid breeds on at least two of the small islets at the southern end of 
the main island; the nesting grounds were discovered by Hubbs. Crossin (MS) estimated 
the Islote Negro population at 800 birds (300 non-breeding) and the Islote Afuera 
population at 4000 birds (1000 non-breeding) in June 1968. In 1977 remains of nine 
birds were found in caves along cliffs at the east side of the island (Pierson and Riedman 
MS), which suggests the possibility of a mainland breeding locale. 

The species occurs near the islands from late December through August, and many 
fly aboard ships at night. The birds apparently first visit the nesting grounds in February. 
Hubbs found none on the islands between October and January (five trips total) but 
found fresh eggs as early as 5 March. The peak of the breeding season is late April- 
June. Yet, the breeding season may be protracted, as Hubbs found fresh eggs as late 
as 29-30 August. Brownell (MS) reported that "adequate nesting grounds on the small 
islotes off Guadalupe are almost fully utilized." If so, nest site limitation would be a 
strong selective agent for an expanded breeding season, as has apparently occurred in 
Leach's Storm-Petrels. 


Geographic variation in the species has been discussed by Jehl and Bond (1975); 
the local form is S. h. hypoleuca. 

Cassin's Auklet {Ptychoramphus aleuticus) 

Although many ornithologists have noted this species at Guadalupe, particularly 
near the southern end of the island (e.g., Thayer and Bangs 1908), it remained for 
Hubbs to discover the nesting area on Islote Negro. The species is not known to nest 
on Islote Afuera (Crossin MS). Brownell (MS) estimated the population at 200 pairs 
in April 1968, a figure that is supported by Hubbs' and Jehl's data. 

Hubbs and associates banded many birds on I. Negro. In 1968 Brownell banded 
56 and recovered two that had been banded two years earlier, one as an adult and one 
as a downy chick. Other banded birds were recovered in April 1970 (Jehl pers. obs.), 
but details are not available. 

The breeding season begins in January. Hubbs reported nests with fresh eggs on 
30 January. The peak in laying occurs by April and by late April many nests may 
contain young. There is annual variation in the nesting period. For example, on 19 
April 1957 Hubbs reported numerous burrows, fresh eggs, eggs with embryos, newly 
hatched young, and well-developed young; on 23 April 1963, 27 nests contained only 
downy young. In most years nesting is completed by late June. On 1 3 June 1955 Hubbs 
found young ready to fledge. On 22 June 1968, most of the colony had completed 
nesting; 85 adults and 40 chicks were present (Crossin MS). 

Rhinoceros Auklet {Cerorhinca monocerata) 

Probably an irregular winter visitor. There are records for 1 9 April 1 925 (McLellan 
1926), 9 February 1957 (Hubbs, specimen LACMNH), and 4 January 1980 (Pitman). 

Rock Dove (Columba livid) 

According to Hubbs, the species was introduced to the island in 1956 by residents 
of the settlement. Kenyon (MS) reported 20 at the weather station in 1965. In 1977 
flocks of up to six were recorded at Twin Canyons and the Lobster Camp (Pierson and 
Riedman MS). Six were seen in the village (along with a peafowl [Pavo cristatus]), on 
5 May 1980 (Everett), and Duncan reported 15-20 there on 19-26 August 1981. 

White- winged Dove {Zenaida asiaticd) 

A specimen of Z. a. mearnsi was collected on 10 June 1953 (Howell and Cade 

Mourning Dove {Zenaida macroura) 

This species was considered accidental by Howell and Cade (1954), perhaps based 
on the report of Gaylord (1897). It has since colonized the island. Hubbs made the 
following observations: 31 August 1956—1; 23 November 1964—1; 10-14 February 
1967 — 24 near the Lobster Camp. By 1970 the species was widespread. Jehl found a 
nest with two young near the village on 14 April and found a pair, almost certainly 
with a nest, at the Lobster Camp on 16 April. Another pair was present on Islote Negro 
on 18 April. In November-December 1973, Mirsky (MS) reported a few at Northeast 
Anchorage and 30 or more near springs. In August 1981, D. Duncan {pers. comm.) 
estimated the population to be in the low hundreds. 

Great Horned Owl {Bubo virginianus) 

The presence of large owls has not been verified. Bryant (1 887a) reported that "Dr. 
Palmer's assistant" stated that a large owl {Bubo) was present on the island, and further 
noted that the Mexican inhabitants reported hearing "hooting" at night. They said, 
however, that the owl was very rare. Ridgway (1876) also noted that "two kinds of 


owls were seen" by the Palmer parly but that no specimens were taken. In 1981, the 
base commander told D. Duncan of large owls in the canyons to the south of the 

Burrowing Owl {Athene cuniculaha) 

This small owl is widespread and common on the main island; it also occurs on 
Islote Negro. The island population is indistinguishable from the mainland form {A. 
c. hypugaea) (Thayer and Bangs 1908). 

Vaux's Swift {Chaetura vau.xi) 

One was seen at the Sealer's Camp on 5 May 1980 (Everett). 

White-throated Swift {Aeronautes saxatalis) 

"Regular visitor, at least formerly. Unreported since 1922" (Howell and Cade 

Anna's Hummingbird {Calypte anna) 

According to Howell and Cade ( 1 954) this hummingbird was evidently uncommon 
to rare prior to 1953 but shortly thereafter became established in the Nicotiana grove 
at Northeast Anchorage. However, Bryant (1887a) was told that they were common 
in palms on the northwestern slope and collected one. Howell and Cade ( 1 954) estimated 
the population at 1 5-20 birds and called attention to the different song of the local 
population, a difference subsequently established by Mirsky (1976). Mirsky estimated 
the population at approximately 100 individuals. On 19 May 1971, Jehl found a nest 
with two eggs in a low shrub near the top of the island, in a canyon above Barracks 

Allen's Hummingbird (Selasphorus sasin) 

Power (1972) incorrectly listed this species as breeding. We know of no evidence 
for its occurrence. 

Belted Kingfisher (Ceryle alcyori) 

Although not listed by Howell and Cade (1954), the kingfisher is an uncommon 
but regular winter visitor. Between 1957-1969, Hubbs had eight records (nine indi- 
viduals) between 25 October and 20 April. It has since been reported almost annually 
(many observers). 

Northern Flicker {Colaptes auratus rufipileus) 

This endemic race was formerly resident in the forested areas atop the island but 
is now probably extinct. Habitat depletion and predation by cats have been considered 
the responsible agents. Apparently it was fairly common and as late as 1906, when last 
seen, the population was reported as "not more than forty individuals" (Thayer and 
Bangs 1908). The history of this local population has been reviewed by Greenway 
(1967); see also Grinnell (1928). 

There are recent reports of flickers at Guadalupe. K. Briggs {pers. comm.) reported 
the species in the pine forest on 17-18 December 1972, and Mirsky (MS) saw one at 
Northeast Anchorage in late November-early December 1973; whether these are fall 
migrants from the mainland or remnants of the endemic population is unresolved. 

Least Flycatcher {Empidonax minimus) 

Accidental. A specimen of this eastern species was taken on 25 October 1962 
(Stager, specimen LACMNH). 


Say's Phoebe (Sayornis saya) 

The only report is of eight at Northeast Anchorage in November-December 1973 
(Mirsky MS). 

Northern Rough-winged Swallow {Stelgidopteryx serripennis) 

Single birds were seen on 23 November 1964 (Hubbs) and on 18 May 1971 (Jehl). 

Barn Swallow {Hirundo rustled) 

Two seen on 19 May 1971 (Jehl) were presumed to be migrants. 

Clark's Nutcracker {Nucifraga columbiand) 

In the invasion year of 1972, at least one nutcracker was observed in the pine 
forest at the north end of the island on 17-18 December (K. Briggs pers. comm.). 

Red -breasted Nuthatch {Sitta canadensis) 

This nuthatch is resident in small numbers in the pine woods at the north end of 
the island. In 1971 Jehl found five pairs there, and on 12-13 April 1970 he observed 
two pairs feeding young and found an additional nest. It occurred in the cypress grove 
in 1953 (Howell and Cade 1954) and probably still does. 

Rock Wren {Salpinctes obsoletus guadalupensis) 

This endemic race is abundant in all open areas of the island, from the beach to 
the crest; it is much less common in forested areas. In 1981, in one open area on top 
of the island, D. Duncan counted one wren per 50 m in a 20 m wide transect. There 
are no current estimates of numbers, but the total population is certainly in the thou- 

Bewick's Wren ( Thryomanes bewickii brevicauda) 

Extinct, last seen in 1892 (Anthony 1901). The history of this endemic form has 
been reviewed by Grinnell (1928) and Greenway (1967). It resided in brushy areas and 
pines, but was never numerous. Habitat depletion by goats and predation by cats 
caused its demise. 

Ruby-crowned Kinglet {Regulus calendula obscurus) 

This endemic race formerly nested in the cypress grove as well as in the pine forest, 
and apparently was fairly common. Howell and Cade (1954) reported five singing males 
in the cypress grove on 1 1 June 1953. Mirsky (MS) reported five in the cypress grove, 
two in the pine-oak grove, and one in the Nicotiana (presumably near the beach) in 
November-December 1973. However, birds seen in winter could be migrants and the 
current status of the endemic population requires verification. 

Mountain Bluebird (Sialia currucoides) 

Three wintered on Guadalupe in 1885-86; one was collected (Bryant 1887a). 

Townsend's Solitaire {Myadestes townsendi) 

One seen on 22 March 1897 (Kaeding 1905) is the only record. 

Hermit Thrush (Cat hams guttatus) 

Bryant (1887a) collected three in the cypress woods between December 1885 and 
March 1886. The race has not been verified (Miller et al. 1957). 


American Robin {Turdus migratorius) 

Bryant (1887a) saw several in December-January 1886-87 in the cypress grove. 
Mirksy (MS) saw one at Northeast Anchorage on 5 December 1973. 

Varied Thrush {Ixoreus naevius) 

One was observed in the pine forest on 4 March 1886 (Bryant 1887^3). 

Northern Mockingbird {Mimus polyglottos) 

Considered accidental by Howell and Cade (1954), apparently on the basis of a 
report by Bryant (1887a), who saw two and collected one on 16 March 1886. One was 
described to Jehl on 22 February 1969. 

Sage Thrasher {Oreoscoptes montanus) 

One was collected on 7 January 1886 (Bryant 1887a). 

Water Pipit {Anthus spinoletta) 

This pipit is probably rare but regular in migration. Bryant reported a flock of 25 
on 2 February 1886 (Bryant 1887a). 

Cedar Waxwing {Bombycilla cedwrum) 

Bryant (1887a) collected one in the winter of 1885-86. 

Loggerhead Shrike {Lanius ludovicianus) 

Bryant (1887a) saw two and collected a female that had fed on a Ruby-crowned 
Kinglet on 29 December 1885. The other bird was heard singing, which suggests the 
possibility of a mated pair. 

European Starling {Sturnus vulgaris) 

On 15 May 1971, Jehl saw one at the settlement at the south end of the island, 
and on 18 May found three in the pine forest at the north end of the island. 

Yellow-rumped (Audubon's) Warbler (Dendroica coronatd) 

This warbler is probably a regular winter visitor. It was first reported by Bryant 
(1887a) and has been seen by many observers. Mirsky (MS) reported up to 30 in 
November-December 1973 at Northeast Anchorage. Everett saw an example of the 
eastern race {D. c. coronatd) on 18 April 1979. 

Townsend's Warbler {Dendroica townsendi) 

Mirsky (MS) reported three in the pine-oak woods on 6 December 1975. 

Black-and-white Warbler {Mniotilta varia) 

One in the pine forest, 19 May 1971 (Jehl). 

Ovenbird {Seiurus aurocapillus) 

A specimen of 5". a. aurocapillus was collected on 9 June 1953 (Howell and Cade 
1954). The late date is typical for eastern vagrants on the west coast in spring. 

Common Yellowthroat {Geothlypis trichas) 

One was collected on 12 November 1938 (Huey 1954). 


Wilson's Warbler ( Wilsonia pusiUa) 

Probably regular in migration but there are only two records; 18 May 1971 (Jehl) 
and 18 April 1979 (Everett). 

Summer Tanager (Piranga rubra) 

A specimen of the eastern race (P. r. rubra) was collected in the cypress grove on 
12 October 1913 (Kimball 1922). 

Rose-breasted Grosbeak {Pheucticus ludovicanus) 

Two records, 24 October 1962 (Stager, specimen LACMNH) and 5 December 
1973 (Mirsky, specimen UCLA). 

Black-headed Grosbeak {Pheucticus melanocephalus) 

The wing of a male was found on the east side of the island on 29 June 1977 
(Pierson and Riedman MS). 

Guadalupe Rufous-sided Towhee (Pipilo erythrophthalmus consobrinus) 

Extinct. The history of this endemic race has been summarized by Grinnell (1928) 
and more fully by Greenway (1967). It was known to occur in the cypress grove and 
perhaps elsewhere, and was last observed in 1897. Its extinction was due to habitat 
depletion by goats and predation by cats. 

Chipping Sparrow (Spizella passerina) 

Bryant (1887<3) collected one on 6 January 1886, and Mirsky (MS) reported the 
species in the Nicotiana and in the pine-oak forest in November-December 1973. 

Fox Sparrow {Passer ella iliaca) 

An example of P. /. sinuousa collected on 16 February 1886 (Bryant 1887a), seems 
to represent the southernmost record for the species on the Pacific coast. 

Lincoln's Sparrow {Melospiza lincolnii) 

Bryant (1887a) collected individuals on 5 and 1 9 February 1886, and Swarth (1933) 
reported a specimen taken on 16 March 1932. 

White-throated Sparrow {Zonotrichia albicollis) 

One collected, 10 October 1913 (Kimball 1922). 

Golden-crowned Sparrow {Zonotrichia atricapilla) 

Bryant (1887a) collected two on 16 February and one on 4 March 1886, in the 

White-crowned Sparrow {Zonotrichia leucophrys) 

Probably regular in migration but the only report is of two near the south end of 
the island on 14 April 1970 (Jehl). 

Guadalupe Dark-eyed Junco {Junco hyemalis insularis) 

Knowledge of this endemic junco was fully summarized by Howell (1968); addi- 
tional information, including variation in the song, was provided by Mirsky (1976). 
At one time the junco was one of the most abundant birds on the island (Palmer, in 
Ridgway 1876). Today it is uncommon and is much less abundant than the House 


Finch or Rock Wren. It may be found scattered along the northern half of the island 
wherever there is vegetation. It often feeds on the ground, in litter at the base of pine 
trees but also in the oaks. However, it seems adaptable and now occupies stands of 
Nicotiana on the beach. Breeding occurs from late January (Bryant 1887a) to at least 
late April (Howell 1 968). On 1 7 May 1 97 1 Jehl saw young juncos that were independent 
of the parents. The taxonomic relationships of this junco have been fully discussed by 
Miller (1941), who argued that the local population was derived from migratory ances- 
tors. "The Guadalupe junco is distinguished principally by its relatively long bill and 
short wing and tail . . . and virtual absence of sexual dimorphism in color" (Howell 
1968). The long bill is used to extract seeds from deep in pine cones (Jehl pers. obs.). 
Power (1980) also discussed the morphology of this species. Bryant (1887a) collected 
a migrant of one of the mainland races {thurberil, cf. Miller et al. 1957) on 6 January 
1886, that was being attacked by a resident junco. 

Western Meadowlark {Sturnella neglecta) 

Bryant (1887a) reported one on the crest of the island on 22 March 1886. 

Brewer's Blackbird {Euphagus cyanocephalus) 

A female was seen on 12 December 1973 (Mirsky MS). 

Scott's Oriole {Icterus parisorum) 

Mirsky (MS) reported two males and three females in the Nicotiana at Northeast 
Anchorage from 23 November to 3 December 1973. 

Guadalupe House Finch {Carpodacus mexicanus amplus) 

This endemic, the second commonest landbird, may occur almost anywhere, in- 
cluding Islote Negro, but is most common near vegetation and at the village. The entire 
population may exceed 1000. Bryant (1887a) provided information on nests and nest 
sites. He also noted that the finches were captured and eaten by locals. The evolution 
and geographic variation of this race have been reviewed by Power (1979). 

Red Crossbill {Loxia curvirostra) 

Howell and Cade (1954) reported this species as "formerly resident; no definite 
breeding record; unreported since 1 903." Evidently it was once fairly common as Bryant 
( 1 887a) reported about 20 in the pines in 1886, and reported collecting nine specimens, 
including an immature in February-March 1896. According to K. C. Parkes, six birds 
collected by A. W. Anthony on 20 September 1896 include a female almost molted 
out of juvenile plumage and five full-grown juveniles. The species was also reported 
as being "resident" by Gaylord (1897), though he did not observe it. Kaeding (1905) 
reported "a few" in 1897. Grinnell (1928) reviewed the status of the species and 
examined the specimens, which he attributed to L. c. bendirei. A. R. Phillips, however, 
now refers all specimens to L. c. benti {fide K. C. Parkes). 

Goldfinch {Carduelis sp.) 

Townsend (1916) states "the Goldfinch was observed." There is no additional 


The Guadalupe avifauna was well-studied in the late 1 9th and early 20th centuries. 
Recent studies have provided new information on seabirds; yet, much remains to be 
learned. The known colonies are diflftcult to reach and most visits to them have been 
made in winter or spring. Studies during other seasons are needed to clarify breeding 


seasons. Efforts are also needed to locate seabird colonies on the main island, especially 
in light of recent reports of storm-petrels calling there at night, and to determine the 
morphological characters of any such populations. It is not inconceivable that the 
Guadalupe Storm-Petrel has escaped extinction. 

The island's rugged topography and lack of fresh water have inhibited recent work 
along the central axis. Surveys in forested areas during the breeding season are needed 
to determine the status of the endemic races of the Ruby-crowned Kinglet and Northern 
Flicker. Both species are common on the mainland and highly migratory, so sight 
records are not proof of the persistence of endemic races. Even evidence of breeding 
may be equivocal, as secondary invasions by these species could have taken place. 
These studies will be difficult and will require capturing or collecting some birds. 

Faunas of oceanic islands are not constant. New species arrive regularly; some 
become established, and others disappear. The factors that affect successful colonization 
or promote extinction are difficult to establish (Jehl and Parkes 1983) but are critical 
to understanding avian distribution. On Guadalupe, a new food source {Nicotiana 
glauca), may have been a major factor in allowing Anna's Hummingbirds to colonize 
(or become more common?) in the past several decades. Mourning Doves have also 
become established, though the reasons why are unstudied. 

In view of the importance of island faunas to current theories in biogeography, regular 
surveys should be encouraged (e.g., at least every decade) so that changes can be detected 
as they are occurring or shortly afterward. Such data will be especially useful from 
islands, like Guadalupe, where a strong historical record has been established. 


We are indebted to Laura C. Hubbs and Elizabeth N. Shor for access to the field 
notes of the late Carl L. Hubbs, and to the following for permitting us to include their 
unpublished records: M. Bonnell, K. Briggs, R. Brownell, E. Chu, R. Condit, R. Crossin, 
R. DeLong, D. Duncan, K. Kenyon, B. LeBoeuf, E. Mirsky, M. Pierson, R. Pitman, 
M. Riedman, B. Tyler, and R. Wisner. 

R. DeLong, R. McConnaughey and R. Moran assisted in various aspects of the 
field research. We especially acknowledge the kindness of the late Carl L. Hubbs, who 
made it possible for Jehl and many others to participate in research at Guadalupe. 

The photograph of the Guadalupe Storm-Petrel was provided by the Field Museum 
of Natural History through the courtesy of M. Traylor. K. C. Parkes, G. Pregill, and 
D. Steadman made helpful comments on an earlier draft of this paper. 

Literature Cited 

Abbott, C. G. 1933. Closing history of the Gua- 
dalupe Caracara. Condor 35:10-14. 

Ainley, D. G. 1980. Geographic variation in 
Leach's Storm-Petrel. Auk 97:837-853. 

. 1983. Further notes on variation in Leach's 

Storm-Petrel. Auk 100:230-233. 

American Ornithologists' Union. 1957. Check- 
list of North American Birds. Fifth Edition. 
Lord Baltimore Press, Baltimore, Maryland. 

. 1983. Check-listofNorth American Birds. 

Sixth Edition. Allen Press, Lawrence, Kansas. 

Anon. 1 874. Guadalupe. La isla de la piel de oro, 
sin duda alguna. Forest and Stream 2(22):337- 

Anthony, A. W. 1898a. Two new birds from the 
Pacific coast of America. Auk 15:36-38. 

. 1898/?. Four sea birds new to the fauna 

of North America. Auk 15:38-39. 

. 1 898c. Petrels of Southern California. Auk 


. 1900. Nesting habits of the Pacific Coast 

species of the genus Puffinus. Auk 1 7:247-252. 


1901. The Guadalupe Wren. Condor 

. 1925. Expedition to Guadalupe Island, 

Mexico, in 1922. The birds and mammals. 
Proceedings California Academy Sciences, Se- 
ries 4 14:277-320. 

Austin, O. L., Jr. 1952. Notes on some petrels of 
the North Pacific. Bulletin Museum Compar- 
ative Zoology. 107:391-403. 

Bent, A. C. 1922. Life histories of North Amer- 
ican petrels and pelicans and their allies. United 
States National Museum Bulletin 121. 

Blake, S. F. 1961. Edward Palmer's visit to Gua- 
dalupe Island, Mexico, in 1875. Madrono 16: 

Bond, J., and R. Meyer deSchauensee. 1944. Fifth 
George Vanderbilt Expedition (1941). Acade- 
my Natural Sciences Philadelphia Mono- 
graph 6. 

Bond, R. M. 1943. Variation in western sparrow 
hawks. Condor 45:168-185. 

Bourne, W. R. P., and J. R. Jehl, Jr. 1982. Vari- 


ation and nomenclature of Leach's Storm-Pe- 
trel. Auk 99:1 92,-1 91 . 

Brown, L., and D. Amadon. 1968. Eagles, Hawks 
and Falcons of the World. 2 vols. McGraw- 
Hill, New York. 

Brownell, R. L., Jr. MS. Preliminary report east- 
em area cruise number 40, Isla Guadalupe 
[1968]. Smithsonian Institution, Pacific Ocean 
Biological Survey Program. 8 pp. 

Bryant, W.E. 1887a. Additions to the ornithology 
of Guadalupe Island. Bulletin California Acad- 
emy Sciences Series 2 2:269-318. 

. 1887ft. Description of a new subspecies 

of petrel from Guadalupe Island. Bulletin Cal- 
ifornia Academy Sciences, Series 2 2:450-451. 
1889. A catalogue of the birds of lower 

California, Mexico. Proceedings California 
Academy Sciences, Series 2 2:237-320. 

Chu, E. W., B. Tyler and D. Lewis. MS. Avian 
notes, pp. 12-31 in M. Pierson (ed.). Report 
of a Scripps Institution of Oceanography Ex- 
pedition to Baja California Islands, 9-20 Feb. 

Connors, P. G. 1983. Taxonomy, distribution, 
and evolution of Golden Plovers Pluvialis do- 
minica and Pluvialis fulva. Auk 100:607-620. 

Crossin, R. S. MS. Preliminary report of Gua- 
dalupe Island [1968]. Smithsonian Institu- 
tion, Pacific Ocean Biological Survey Program. 
8 pp. 

. 1974. The slorm-peXTeh (Hydrobatidae). 

pp. 154-205 />2 W. B. King(ed.). Pelagic studies 
of seabirds in the Central and Eastern Pacific 
Ocean. Smithsonian Contributions Zoology 

Davidson, M. E. McLellan. 1928. On the present 
status of the Guadalupe petrel. Condor 30:355- 

DeLong, R. L. and R. S. Crossin. MS. Status of 
seabirds on Islas de Guadalupe, Natividad, 
Cedros, San Benitos, and Los Coronados. 

Devillers, P., G. McCaskie. and J. R. Jehl, Jr. 1971. 
The distribution of certain large gulls {Lams) 
in southern California and Baja California. 
California Birds 2:11-26. 

duPetit-Thouars, A. 1956. Voyage of the Venus 
Sojourn in California. Excerpt from "Voyage 
autour du monde sur la fregate Venus pendant 
les annees 1836-1839. Translated by C. N. 
Rudkin. Los Angeles, Glen Dawson Press. 

Emerson, W. O. 1906. Oceanodroma leucorhoa 
and Its relatives on the Pacific coast. Condor 

Friedmann, H., L. Griscom, and R. Moore. 1950. 
Distributional check-list of the birds of Mex- 
ico. Pacific Coast Avifauna 29. 

Gaylord, H. A. 1897. Notes from Guadalupe Is- 
land. Nidologist 4:41-43. 

Godman, F. D. 1907-1910. A monograph of the 
petrels. Witherby, London. 

Greenway, J. C. 1967. Extinct and Vanishing Birds 
of the New World. Dover Publications. New 

Grinnell,J. 1918. The status of the white-rumped 
petrels of the California coast. Condor 20:46. 

. 1928. A distributional summation of the 

ornithology of Lower California. University 
California Publications Zoology 32(1): 1-300. 

Hanna, G Dallas. 1925. Expedition to Guadalupe 
Island, Mexico, in 1922. General Report. Pro- 
ceedings California Academy Sciences Series 4 

Howell, T.R. 1968. Guadalupe Junco. pp. 1094- 
1098 m A. C. Bent et al. (O. L. Austin, Jr. 
[ed.]). Life histories of North American car- 
dinals, grosbeaks, buntings, towhees, finches, 
sparrows, and allies. United Stales National 
Museum Bulletin 237. 

, and T. J. Cade. 1954. The birds of Gua- 
dalupe Island in 1953. Condor 56:283-294. 

, and . 1955. Additional data on the 

birds of Guadalupe Island. Condor 58:78. 
Hubbs, C. L. 1960. The marine vertebrates of the 

outer coast. Systematic Zoology 9:134-147. 
, and J. R. Jehl, Jr. 1976. Remains of Pleis- 
tocene birds from Isla de Guadalupe, Mexico. 

Condor 78:421-422. 
Huey, L. M. 1924. A trip to Guadalupe, the isle 

of my boyhood dreams. Natural History 24: 

. 1925. Guadalupe Island: an object lesson 

in man-caused devastation. Science 61:405- 

. 1930. Notes on the habits and plumage 

of young Kaeding Petrels. Condor 32:68-69. 
. 1952. Oceanodroma tethys tethys, a petrel 

new to the North American avifauna. Auk 69: 

460-46 1 . 
. 1954. Notes from Southern Cahfomia and 

Baja California, Mexico. Condor 56:51-52. 

Jehl, J. R., Jr. 1972. On the cold trail of an extinct 
petrel. Pacific Discovery 25(6):24-29. 

, and S. I. Bond. 1975. Morphological vari- 
ation and species limits in murrelets of the ge- 
nus Endomychura. Transactions San Diego 
Society Natural History 18:9-23. 
-, and K. C. Parkes. 1983. "Replacements" 

of landbird species on Socorro Island, Mexico. 
Auk 100:551-559. 

Johnson, C. W. 1953. Notes on the geology of 
Guadalupe Island, Mexico. American Journal 
Science 251:231-236. 

Kaeding, H. B. 1905. Birds from the west coast 
of Lower California and adjacent islands. Con- 
dor 7:105-138. 

Kenyon, K. W. MS. Expedition to Baja Califor- 
nia, Mexico 19 Jan.-l 1 Feb. 1965. Unpub- 
lished report to U.S. Fish and Wildlife Service. 

Kimball, H. H. 1922. Bird records from Califor- 
nia, Arizona, and Guadalupe Island. Condor 

Lewis, L. R. 1971. Baja Sea Guide, Vol. 2. Miller 
Freeman Publications, San Francisco. 

Lindsay, G. 1966. Guadalupe Island. Pacific Dis- 
covery 19:2-1 1. 

Loomis, L. M. 1918. Expedition of the California 
Academy of Sciences to the Galapagos Islands, 
1905-1906. 12. A review of the albatrosses, 
petrels, and diving petrels. Proceedings Cali- 
fornia Academy Sciences Series 4 2:1-187. 

Madden, H. M. 1949. Xantus, Hungarian natu- 
ralist in the pioneer west. Palo Alto, California. 
Books of the West. 

McLellan, M. E. 1926. Expedition to the Revil- 
lagigedo Islands, Mexico, in 1925. VI. The birds 


and mammals. Proceedings California Acad- 
emy Sciences Series 4 15:279-322. 

Miller, A. H. 1941. Speciation in the avian genus 
Junco. University California Publications Zo- 
ology 44:173-434. 

Miller, A. H., H. Friedmann, L. Griscom, and R. 
T.Moore. 1957. Distributional check-list of 
the birds of Mexico. Pacific Coast Avifauna 
No. 33. 

Mirsky, E. N. MS. The Guadalupe Island avifau- 
na in 1973. 

. 1976. Song divergence in hummingbird 

and junco populations on Guadalupe Island. 
Condor 78:230-235. 

Oberholser, H. C. 1917. A review of the subspe- 
cies of the Leach Petrel, Oceanodroma leucor- 
hoa (Vieillot). Proceedings United States Na- 
tional Museum 54:165-172. 

Palmer, R. S. (ed.). 1962. Handbook of North 
American Birds. Vol. 1 . Yale University Press, 
New Haven and London. 

Pierson, M., and M. Riedman MS. Report of an 
expedition to Isla de Guadalupe, Baja Califor- 
nia, Mexico 20 June-13 July 1977. Unpub- 
lished Report, Univ. Calif. Santa Cruz. 1 5 pp. 

Power, D. M. 1972. Numbers of bird species on 
the California Islands. Evolution 26:451-463. 

. 1979. Evolution in peripheral isolated 

populations: Carpodacus finches on the Cali- 
fornia Islands. Evolution 33:834-847. 

. 1980. Evolution of land birds on the Cal- 
ifornia Islands, pp. 613-650 in D. M. Power 
(ed.). The California Islands: Proceedings of a 
multidisciplinary symposium. Santa Barbara 
Museum Natural History. 
-, and D. G. Ainley. MS. Similarity among 

populations of Leach's Storm-Petrels. 
Ridgway, R. 1876. Ornithology of Guadeloupe 
[sic] Island, based on notes and collections made 
by Dr. Edward Palmer. United States Geolog- 
ical and Geographical Survey Territories 2:183- 

Rothschild, W., and E. Hartert. 1902. Further 
notes on the fauna of the Galapagos Islands. 
Notes on birds. Novitates Zoologica 9:381- 

Small, A. 1961. Southern Pacific Coast Region. 
Audubon Field Notes. 15:438. . 

Swarth, H. S. 1933. Off-shore migrants over the 
Pacific. Condor 35:40. 

Thayer, J. E., and O. Bangs. 1908. The present 
state of the omis of Guadaloupe (sic) Island. 
Condor 3:101-106. 

Thobum, W. W. 1899. The birds of Guadalupe 
Island, p. 278 in D. S. Jordan. The fur seals 
and fur-seal islands of the North Pacific Ocean. 
Pt. 3. United States Government Printing Of- 
fice, Washington, D.C. 

Todd, W. E. C. 1955. Taxonomic comment on 
races of Leach Petrel of the Pacific Coast. Con- 
dor 57:122. 

Townsend, C. H. 1890. Scientific results of ex- 
plorations by the U.S. Fish Commission 
Steamer Albatross, No. XIV. — Birds from the 
coasts of western North America and adjacent 
islands, collected in 1888-89, with descrip- 
tions of new species. Proceedings United States 
National Museum 13:131-142. 

. 1911. (Account of expedition to lower 

California). Auk 28:390-391. 

. 1916. Voyage of the 'Albatross' to the 

Gulf of California in 1911. Bulletin American 
Museum Natural History 35:399-476. 

. 1923. Birds collected in Lower California. 

Bulletin America Museum Natural History 48: 

vanRossem, A. J. 1942. Preliminary comment on 
some Pacific Coast petrels. Proceedings Bio- 
logical Society Washington 5:9-12. 

von Berlepsch, H. G. 1906. On a new form of 
Oceanodroma inhabiting San Benito Island, off 
the coast of Lower California. Auk 13:185- 


2044 093 362 671