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Chapter 64. Evolution and biogeography of Orionina in the Atlantic, Pacific and Caribbean : Evolution and speciation in Ostracoda, II

Chapter 64. Evolution and biogeography of Orionina in the Atlantic, Pacific and Caribbean : Evolution and speciation in Ostracoda, II

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928 T. M.



CRONIN AND



N. SCHMIDT



After its original description based on species from Florida by Puri (1953), Bold (1963) redescribed Orionina in detail, giving a comprehensive list of occurrences from the Caribbean and

Central America. The type species, 0. vaughani was described in 1904 by Ulrich and Bassler based

on material from the Pliocene of Virginia. Although our results and interpretations of some of

the relationships among species differ slightly from those of van den Bold, his work stands as the

underlying framework for all subsequent study of this genus. Gunther and Swain (1976) provided

important occurrence data from the Gulf of Panama and postulated some evolutionary

relationships in Orionina. Other important occurrences of Orionina from the eastern Pacific are

found in Swain (1967, 1969), Swain and Gilby (1967), and Valentine (1976). Holden (1976)

described Orionina flabellacosta from the Miocene of Midway Island, Hawaii. A new species

closely related to Holden's species is proposed below.



LENGTH vs. HEIGHT in OR/O/U/NA

0

400



380



-



38C



c



e



.o



E

Y



+



340



320



(3



i



I



300



280



260



240



220



420



~~



480



600



640



680



820



880



700



740



780



820



TEXT-FIG.

1-Plot of carapace length versus height for Orioninu from various localities. The vuughuni group,



designated by squares, includes specimens assigned by other authors to 0. pseudovuughani and 0 .

serruluta. Holotype specimens are indicated by open stars (measurements of 0. brudyi, 0. sirnilis, 0.

serrulatu, and 0. vuughuni holotypes are taken from Bold, 1963).



880



Evofution and Biogeography of Orionina 929



LENGTH vs. RIDGE ANGLE in OR/ON/NA

84



0

0



.



0



8:



00



0



ec



o



OOIOD.~



0



H



m



m



rn



v



D



RIDGE ANGLE



7f



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j



74



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0



n

ti



70



88



86



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m

82



.



o



4



60



4



4



56



860



LENGTH (microns)

TEXT-FIo.2-Plot of carapace length versus angle between median longitudinal ridge and oblique ridge in

posterior part of the carapace. See Text-fig. 1 for explanation of symbols.



MATERIAL

AND METHODS

Table 1 lists the formations and recent material from which specimens of Orionina were exPoag. 1972 and 0. butlerae Bold. 1965 were not examined, nor was 0.fragilis Bold, 1963. One hundred and sixty one specimens from our collection

amined Pre-late Miocene snecies 0.armata



._.^_

-I-^* ----- L - 1



.



--.:*l-



,,,,,:..rr



,1,&,-.



,:,.,-,.,



..A



..":..".,



A:":+:"-..



...nno..mmn~4n



.I,P..P



taken of carapace length, anterior and posterior height, and distance from the anterior margin to

the muscle platform. We also measured the angle between the median longitudinal ridge and the

oblique ridge (see Text-fig. 2). These data were subjectedto various bivariate analyses of which two

plots are presented here.



INTERSPECIFIC

VARIATION

Table 2 lists the fourteen proposed species of Orionina and the taxonomic assignment given in

the present paper. We recognize ten species including the three unstudied species 0. armata, 0.



930 T. M. CRONINAND N. SCHMIDT

TABLE1-Orionina MATERIAL

STUDIED.

t

MIOCENE

Formation

Manzanilla

Gatun

Cubagua

Cueva de Angostura

Undescribed



Region

Trinidad

Panama

Venezuela

Ecuador

Marshall Islands



PLEISTOCENE

Formation

James City

Chowan River

Canepatch

Waccarnaw

Caloosahatchee

BeILmont

Fort Thompson

Undescribed



Region

North Carolina

North Carolina

South Carolina

South Carolina

Florida

Florida

Florida

Marshall Islands



PLIOCENE

Region

Yorktown

Virginia

Duplin

North Carolina

Raysor

South Carolina

Goose Creek

South Carolina

Bear Bluff

South Carolina

Tamiarni

Florida

Pinecrest

Florida

Jackson Bluff

Florida

Bowden

Jamaica

Gurabo

Dominican Republic

Limon

Costa Rica

HOLOCENE

Region

Bermuda

Acklins Island, Bahamas

Florida Keys

St. Thomas, U.S. Virgin Islands

Dominican Republic, North Shore

Dominican Republic, South Shore

Payardi Island, Panama

Gulf of Panama

Campeche Banks, Mexico

Corinto Bay, Nicaragua

Fiji

Formation



t, Exact locality information is available from T. M. Cronin.

TABLE2-DESCRIBED



SPECIES OF OriOnina.



Species

t 0, armata Poag, 1972

Cythere bermudae Brady, 1880

0 . bradyi van den Bold, 1963

0. brouwersae new species

t 0 . butlerae van den Bold, 1965

0. ebanksi Teeter, 1975

0. eruga van den Bold, 1963

0. flabellacosta Holden, 1976

t 0. fragilis van den Bold, 1963

0. boldi new species

0 .pseudovaughani Swain, 1967

Cythereis reticulata Hartmann, 1956

Cythere serrulata Brady, 1869

0 . similis van den Bold, 1963

Cythere vaughani Ulrich and Bassler, I904

t, Not Studied.



This Paper

0 . armata

0 . bradyi

0 . bradyi

0 . brouwersae n. sp.

0 . butlerae

0. ebanksi

0 . eruga

0.flabellacosta

0 .fragilis

0 . boldi n. sp.

0 . vaughani

0. vaughani

0 . vaughani

0 . similis?

0 . vaughani



fragilis, and 0 . butlerae, and the seven studied for this paper-0. boldi, 0. bradyi, 0 .brouwersae,

0 . ebanksi, 0 . eruga, 0 . flabellacosta, and 0 . vaughani. Each species consists of populations

having a distinct morphotype consistently distinguishable from the other species in populations

examined from most of their stratigraphical and geographical ranges. Plate 1 illustrates the seven



Evolution and Biogeography of Orionina 931



species and Text-fig. 1 plots their carapace lengths and heights. These figures should be referred

to in the following summaries of the characteristics of each species.

Orionina vaughani is comprised of a complex of populations all having relatively large carapaces, strong sexual dimorphism, well-developed ridges and surface reticulation, and, in some

populations, variable radial pore patterns. The intraspecific variation and synonymy of 0. vaughani

with other species is discussed below. 0. bradyi has a small carapace, lacks the vertical posterior

ridge present in other species, and has a low angle between the medial and oblique ridges (see P1. 1,

fig. 1). Sexual dimorphism is not as pronounced as in other species of Orionina. 0. eruga displays

a very large, elongate carapace, a strongly convex dorsum and less-tapered posterior end. The

murae forming the reticulum are not as sharp as in 0. vaughani. 0. ebanksi is very similar to 0.

bradyi, especially in its small size and evenly reticulate ornamentation. It differs in having its dorsal

ridge merge gradually with the dorsum in contrast to the sharper intersection in 0. bradyi and in

the curved shape of the dorsal margin. 0. brouwersae is quite distinct from other members of the

genus and is characterized by a very high carapace (Text-fig. 1) and a distinct pattern of longitudinal

ridges. Also, the anterior ridge runs through the eyespot continuing to the dorsal margin in contrast to the ridge position in other species, 0.flabellacosta lacks the prominent medial and vertical

ridges typical of Caribbean and Atlantic Orionina, but has the characteristic curved ventral ridge

and reticulate surface. Gunther and Swain (1976) illustrated a large, strongly calcified species, which

they designated as Orionina n. sp. 2, from the Gulf of Panama. We have found this form in the

Pliocene of Costa Rica, and in modern sediments off the Florida Keys. We describe this new species as 0. boldi below.



INTRASPECIFIC

VARIATION

We gave considerable attention to the relationships between Orionina vaughani and the species

0. serrulata, 0. pseudovaughani, and 0 . reticulata, each long recognized as closely related, but

usually separated on the basis of minor differences. Specimens that have been identified as 0.

serrulata are said to be identical to vaughani in external appearance, differing only in the presence of

indentations in the line of concrescence and bundling of radial pores, whereas 0. vaughani has

numerous parallel, straight pores (Bold, 1963). In a preliminary study of Gulf of California

ostracods, Swain et al. (1964) identified 0. vaughani for the first time from the Pacific, but later

Swain (1967) redescribed these populations as 0. pseudovaughani, which he characterized as

differing from 0. vaughani in having three rather than four pillars in the anterior vestibule. Swain

also pointed out that these populations might be conspecific with 0. serrulata, but he considered

0. serrulata to have a confused taxonomic history.

We regard 0. serrulata, 0.pseudovaughani, and 0 .reticulata as conspecific with 0. vaughani and

believe that the minor differences among them represent interpopulational variation, perhaps

related to local environmental conditions. We don’t consider the variation in radial pores a specific

level character because we have observed populations from the Limon of Costa Rica that have

individuals with both the straight, parallel pattern and the bundled pattern. Similarly, Gunther

and Swain (1976) illustrate these two types of pore pattern in populations from the Gulf of

Panama and they assign them to two species (0.vaughani and 0 .pseudovaughani). It is our opinion

that these two morphotypes belong to a single species. Many other shallow marine genera, such

as Proteoconcha (Plusquellec and Sandberg, 1969; Hazel and Cronin, this volume) show strong

variation in radial pores. Text-fig. 1 shows the similarity in carapace length and height of Orionina

populations from the eastern United States, Central America and the Caribbean, the eastern



932 T. M. CRONIN AND N. SCHMIDT



Pacific and Florida. The SEM photographs in Plate 2 show the similarity in surface ornamentation

of populations from different regions. The evidence shows that intraspecific variation is

substantially less than interspecific variation within the genus. Plate 2 illustrates this point with

four specimens of 0. boldi n. sp. This species almost certainly descended from Caribbean

populations of 0. vaughani but the two species are consistently distinct from one another in all the

populations studied. There are no gradational populations between the two species.



GEOGRAPHICAL

ISOLATION

IN 0RZONINA

The formation of the Isthmus of Panama during the Pliocene, about 3 to 4 million years ago, represents a major barrier to the genetic interchange of eastern Pacific and Caribbean populations

that were formerly contiguous with each other. That Orionina lived on both sides before the Isthmus

developed and was not artifically introduced into the Pacific is demonstrated by Miocene and

early Pliocene occurrences throughout the Caribbean and southeastern North America (Bold,

1963), in the Pacific at Midway Island (Holden, 1976), and reported here for the first time, in

Ecuador and the Marshall Islands. Evidence from the two species known as fossil and/or Holocene

from both sides indicates minor morphological divergence of populations since separation several

million years ago. Orionina vaughani is known pre-and post-Isthmus on both sides and comparison

of specimens illustrated in Plate 2 show a strong morphological similarity between Pacific and

Atlantic/Caribbean forms. 0. bold occurs in Holocene sediments on the Pacific side in the Gulf

of Panama and off southern Florida. As a fossil, it is known from the Pliocene of Costa Rica.

Although much less common than 0. vaughani, there are no consistent differences in carapace

morphology in populations from the different regions (PI. 2).

A second type of barrier isolating shallow water populations is deep water. Deep water separates many populations of Orionina living off Caribbean Islands from shelf populations off the

coasts of the Americas and from each other. However, the distances across these barriers is relatively

small and they were even less during the numerous glacial periods when sea level was 100 to 150

metres lower that its present level. Consequently, there is a much higher probability of passive

dispersal of ostracods (possibly on drifting seaweed, see Teeter, 1973; Cronin, this volume) among

western Atlantic, Caribbean and Gulf of Mexico populations than there is among more distant

islands such as the Hawaiian Islands, the Marshall Islands and off Fiji, where 0. Jabellacosta

and 0. brouwersae occur. In the case of these two Pacific species, alternative hypotheses might

explain the seemingly anomalous occurrence of Orionina, a genus otherwise restricted to the

eastern Pacific, western Atlantic and Caribbean. One alternative is that these species are not phy-



PLATEl-Fig. 1 . Orionina bradyi Bold, 1963. Lateral view, left valve, female (USNM 403818, Holocene, Bahamas),

x 78 Figs. 2, 4. Orionina brouwersae n. sp. 2. Internal view, left valve, female (USNM 403819, Recent, Fiji),

x 78 4. Lateral view, left valve, female (Holotype, USNM 403820, early Miocene-late Pliocene, Marshall

Islands), x 78 Fig. 3. Orionina cf. 0. brouwersae Cronin and Schmidt n. sp. Lateral view, left valve, female

(USNM 403821, early Miocene-late Pliocene, Marshall Islands), x78 Figs. 5, 6, 9, 10. Orionina vaughani

(Ulrich and Bassler, 1904). 5. Lateral view, left valve, female (USNM 403822, Pliocene, Costa Rica), x 78

6. Lateral view, left valve, male (USNM 403823, Pliocene, Costa Rica), x78 9. Lateral view, left valve,

female (USNM 403824, Pliocene, North Carolina), x 7 8 10. Lateral view, left valve, male (USNM 403825,

Pliocene, North Carolina), x78 Figs. 7, 8. Orionina boldi n. sp. 7. Lateral view, left valve, female

(Holotype, USNM 403826, Holocene, Florida), x 78 8. Lateral view, left valve, male (USNM 403827,

Holocene, Gulf of Panama), x 7 8 Figs. 11, 12. Orionina cf. 0. eruga Bold, 1963. 1 1 . Lateral view, left

valve, female (USNM 403831, Pliocene, Venezuela), x 78 12. Lateral view, left valve, male (USNM 403832,

Pliocene, Venezuela), x 78 Fig. 13. Orionina ebanksi Teeter, 1975. Lateral .view, left valve, female (USNM

403833, Holocene, St. Thomas, U. S. Virgin Islands), x78



Evolution and Biogeography of Orionina 935



logenetically related to Caribbean Orionina, but are descended from another western Pacific hemicytherine lineage. At present, well-documented Miocene to Holocene faunas from Midway (Holden,

1976) and the Marshall Islands (Cronin, unpublished data) do not show any likely candidates.

Further, as recognized by Holden, external and internal carapace features are clearly Orioninalike, although these central and western Pacific forms have evolved so that they are distinct from

other species in details of carapace ornament. We consider the most reasonable interpretation to

be that 0. flabellacosta split from early Miocene populations of Orionina that became isolated

in the Hawaiian Islands, and that during the middle or late Miocene, 0. brouwersae split when populations became dispersed farther west in the Marshall Islands. Although the Pacific data are still

sparse and phylogenetic relationships unclear, the evidence indicates at least two speciation events

occurred when relatively small populations became isolated on small islands, probably through

passive dispersal on seaweed (Teeter, 1973). These speciation events contrast with the lack of speciation or even intraspecific morphologic divergence that resulted from isolation by the Isthmus of

Panama.

The patterns observed in Orionina are, of course, subject to different interpretations. For example,

Gunther and Swain (1 976) hypothesized that major barriers existed between Jamaica and Florida

and between Venezuela and the eastern Pacific such that “three more-or-less isolated evolving

groups” (p. 141) of Orionina can be recognized. Based on our distribution data and the inferred

dispersal capabilities of tropical ostracod genera (see also Teeter, 1973), we do not believe there

were effective barriers east of Central America within the Caribbean, Gulf of Mexico, and western

Atlantic regions. Although 0. ebanksi and 0. boldi probably evolved within this region, there is no

direct evidence for geographic isolation having been the cause. Further, considering the 25 million

year history of the genus, surprisingly few species have evolved. Even the land barrier formed by

the Isthmus of Panama has not yet led to divergence within Orionina. We believe the preliminary

evidence from the central and western Pacific shows isolation of small populations led to the formation of at least two new species and the most significant morphological divergence within the

genus. Overall, the results support the hypothesis of Valentine and Jablonski (1 983) that geographical isolation of large populations of benthic invertebrates results in slow rates of morphological

change. Conversely, rapid morphological change accompanying speciation events occurs when

small populations become isolated.



SYSTEMATIC

PALEONTOLOGY

Our study was designed to investigate the effects of types of geographical isolation on the genus

Orionina. A major taxonomic revision is beyond the scope of this paper. Further, the phylogenetical relationships of some species remain unclear, so an abbreviated systematic section is given.

PLATE2-Figs. 1-10. Orionina vaughani (Ulrich and Bawler, 1904). 1 . Lateral view, left valve, (USNM 403834,

Holocene, Gulf of Panama), x 7 8 2. Lateral view, left valve, (USNM 403835, early Pleistocene, North

Carolina), x 78 3. Lateral view, left valve, (USNM 403836, early Pleistocene, North Carolina), X 78 4.

Lateral view, left valve, (USNM 403837, Holocene, Gulf of Panama), x 78 5 . Lateral view, left valve,

(USNM 403838, middle Pleistocene, South Carolina), x 78 6. Lateral view, left valve, (USNM 403839,

early Pliocene, South Carolina), x 78 7. Lateral view, left valve, (USNM 403840, Miocene, Ecuador),

x 78 8. Lateral view, left valve, (USNM 403841, Pliocene, Panama), x 78 9. Lateral view, left valve,

(USNM 403842, middle Pleistocene, Florida), X 78 10. Lateral view, left valve, (USNM 403843, late

Miocene, Venezuela), x 7 8 Figs. 11-14. Orionina boldi n. sp. 11. Lateral view, left valve, (USNM 403828,

Pliocene, Costa Rica), x 78 12. Lateral view, left valve, (USNM 403829, Pleistocene, Florida), X 78.

13. Lateral view, left valve, (USNM 403827, Holocene, Gulf of Panama), x 7 8 14. Lateral view, left

valve, (USNM 403830, Holocene, Florida Keys), X 78.



936 T. M. CRONIN

AND N. SCHMIDT



Van den Bold (1963) discussed early studies of Orionina and should be consulted for Caribbean

occurrences, synonymies, and detailed descriptions of most species. We concentrate here on the

two new species and their distinction from other Orionina. As mentioned above, we did not examine 0. armata, 0.fragilis and 0. butlerae, but we consider at least the first two as valid species

distinct from those illustrated here. Orionina butlerae we believe to be in need of further study.

Genus ORIONINA

Puri, 1954

ORIONINA

BOLDI Cronin and Schmidt n. sp.

(PI. 1, figs. 7, 8; P1. 2, figs. 11-14)

Etymology.-Named for Dr. W. A. van den Bold, Louisiana State University, in honor of his

pioneering work on Orionina.

Types.-Holotype, a female left valve from the Holocene off the Florida Coast, USNM 403826

(PI. 1 , fig. 7). Sample 1554 of Hathaway (1971), 55 m water depth 26”31.1’N, 8Oo01.4’W.

Zl1ustratedspecimens.-USNM 403827 (PI. 1, fig. 8 ; PI. 2, fig. 13); USNM 403828 (PI. 2, fig. 11);

USNM 403829 (PI. 2, fig. 12); USNM 403830 (Pl. 2, fig. 14).

Diagnosis.-Very large and heavily calcified, having very thick longitudinal ridges and two

subvertical ridges in the posterior region that connect the longitudinal ridges.

Description.-Carapace slightly more rectangular and less tapered posteriorly than other Orionina, valve walls very thick and heavily calcified. Lateral surface of valves having two prominent

longitudinal ridges that converge near the muscle node and that diverge anteriorly and posteriorly

from the anterocentral region. Posteriorly, there is a strong nearly vertical ridge and anterior of

this, a distinct oblique ridge connecting the longitudinal ridges. Postero-ventral ridge is sometimes

nodose. Between the ridges the surface is variably reticulate to almost smooth. Internal features

are typical for the genus and there are many straight, parallel, evenly spaced radial pore canals.

Remarks.-Some female specimens resemble large males of 0. vaughani, but differ in their

larger size, and the presence of the well-developed oblique ridge, anterior to the posteroventral

ridge, that connects the longitudinal ridges. Some specimens resembling 0. boldi were found in

the Bowden Formation of Jamica, but they were much smaller and require additional study.

Occurrence.-Recent off South Florida and Gulf of Panama; Bermont Formation, middle

Pleistocene, subsurface of South Florida; Limon Formation, Pliocene, Costa Rica.

Bold, 1963

(PI. 1, fig. 1)



ORIONINA BRADYI



Remarks.-This widespread species is well-known throughout the Caribbean (Bold, 1963). DiasBrito et al. (this volume) has also found it in Sepetiba Bay, Brazil. Bold (1963, p. 45-47) considers

some specimens described by Brady (1880) as Cythere bermudae to be synonymous with 0. bradyi

and discusses the taxonomy of these species in detail.



ORIONINA

BROUWERSAE Cronin and Schmidt n. sp.

(PI. 1, figs. 2,4)

Etymology.-Named for Dr. Elisabeth M. Brouwers, U. S. Geological Survey.

Types.-Holotype, a female carapace, USNM 403820, from the Pliocene (a depth of 498 feet in

a core) from Enewetak Atoll, Marshall Islands (PI. 1, fig. 4).

Illustrated specimens.-USNM 403819 (PI. 1, fig. 2).

Diagnosis.-Relatively low length/height ratio, valve surface evenly reticulate, having two

short parallel anterior ridges, two fine posterior ridges, an anterior marginal ridge running through

the eye tubercle.



Evolution and Biogeography of Orionina 937



Description.-Carapace arched, convex dorsally, evenly rounded anteriorly, surface ornament

consisting of reticulation and polygonal fossae. An anterior marginal ridge runs through eye

tubercle continuing along dorsal margin. Longitudinal ridges differ from those in other Orionina,

consisting of two short anterior ridges, two fine, posterodorsal ridges and a long ventral ridge.

Ridges and murae are relatively thin compared to Caribbean species.

Remarks.-Plate 1, figure 3 shows a morphotype that is tentatively assigned to this species

based on its size shape and reticulate surface. However it clearly differs from the Holotype illustrated in Plate 1, figure 4 in the development and orientation of posterodorsal and to a lesser extent in

the anterocentral ridges. Both these specimens come from the subsurface Miocene-Pliocene of

Enewetak Atoll. Specimens from recent sediments of Fiji are more similar to the holotype specimen and are confidently placed in this species. Orionina flabellacosta Holden 1976, differs from

0. brouwersae in having a curved ventral ridge, lacking the short anterior ridges and having larger

male carapaces.

Occurrence.-Miocene t o Pleistocene of Enewetak Atoll; Recent off Fiji.



ORIONINA

cf. 0. ERUGA Bold, 1963

(Pl. 1, figs. 11, 12)

Remarks.-The specimens illustrated in Plate 1 from the Pliocene Cubagua Formation of

Venezuela have a stronger ridge than those illustrated by Bold (1 963) from the Miocene of Trinidad.



ORIONINA

FLABELLACOSTA Holden, 1976

Orioninaflabellacosta HOLDEN,

1976, p. F20, PI. 3, figs. 5-7; PI. 1 1 , fig. 6.

Remarks.-Holden

Hawaiian Islands.



(1 976) described and illustrated this species from the Miocene of Midway,



ORIONINA

VAUGHANI (Ulrich and Bassler, 1904)

(PI. 1, figs. 5, 6, 9, 10; PI. 2, figs. 1-10)

Cythere serrulata BRADY,1869, p. 153, PI. 18, figs. 1 1 , 12.

Cythere vuughani ULRICH

AND BASSLER,

1904, p. 109, PI. 38, fig. 25.

Cythereis reticulata HARTMANN,

1956, p. 37, figs. 45-52.

Orioninapseudovaughani SWAIN,

1967, p. 86, PI. 3, figs. 5a, b; PI. 4, figs. 6a-c; Text-figs. 50d-g, 5 4 ~ .

Remarks.-As discussed in detail above, we believe Orionina vaughani, 0. serrulata, 0. pseudovaughani and 0. reticulata constitute different populations and morphotypes of the same species.

Although serrulata was described by Brady in 1869, Bold (1963) was not able to find its holotype

and Swain (1967) expressed concern over the confusing status of this species. We therefore use

vaughani to refer to this well known species.



ACKNOWLEDGMENTS

We gratefully acknowledge the help of the following colleagues who freely loaned us specimens

of Orionina for study: Drs. M. L. Gamero, J. E. Hazel, M. Kontrovitz, F. M. Swain, W. A. van

den Bold, T. R. Waller. Thanks go to Lt. Col. Robert F. Couch, Jr., Ms. E. E. Compton-Gooding

and Dr. E. M. Brouwers for their help with the Marshall Islands material. Dr. Joseph E. Hazel

(Louisiana State University) and Dr. Norman F. Soh1 (U. S . Geological Survey) provided helpful

reviews of the manuscript and invaluable taxonomic advice.



AND

938 T. M. CRONIN



N. SCHMIDT



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BOLD, W.A. VAN DEN.



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Chapter 64. Evolution and biogeography of Orionina in the Atlantic, Pacific and Caribbean : Evolution and speciation in Ostracoda, II

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