Contributions to Zoology, 68 (1) 19-35 (1998)

SPB Academic Publishing bv, Amsterdam

A new spelaeogriphacean (Crustacea: ) from the Upper of China

Shen Yan-bin Rod+S. ² & Frederick+R. Schram² ¹, Taylor

1Nanjing Institute of Geology and Palaeontology, Academia Sinica, 39 East Beijing Road, Nanjing, The

2 People’s Republic of China, 210008; Institute for Systematics and Population Biology, University oj

Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands

Keywords: Crustacea, Peracarida, Spelaeogriphacea, , Upper Jurrasic north-east China, cladistic

analysis

Abstract phylogenetic significance for Spelaeogriphus

lepidops by Grindley and Hessler (1971).

A of is described new monotypic Spelaeogriphacea from While work on S. lepidops has been sparse, the Upper Jurassic ofLiaoning Province, north-east China. This continuing research on other fronts has increased new genus and brings the number of known spelaeo- the number of species credited to this order to three. griphacean taxa to four, the others being two recent forms from second Pires described a recent form, Brazil and (1987) South Africa, and one from the of Potiicoara found in a lake inside the eastern Canada. brasiliensis, The new Chinese form is morphologically(and

similar the calcareous cave Gruta do Azul in the Bodo- phylogenetically) very to recent spelaeogriphaceans, Laga

that the seenin the recent do suggesting body plan Spelaeogriphacea quena Mountains, MatoGrosso Sul, Brazil. This

was achieved in the of the A relatively early history group. form is distinguished from S. lepidops by its shorter cladistic analysis of this and several other peracaridan orders biramous fifth carapace, pleopod, three-segmented indicates that the be Spelaeogriphaceamay a paraphyletic group. mandibular endite and This palp, spinose maxillipedal suggests that much work remains to be done with respect collection to the and the its specific locality, phylogenetic relationships among highly considerably that S. peracaridan taxa. distant from of lepidops. Pires also included

a phylogenetic analysis of the peracaridan orders,

considering the phylogeny of the order as a whole. Introduction Of particular note, she proposed as a result of this

analysis a sister-group relationship between the

As a result of several specimens obtained by the Spelaeogriphacea and . South African Spelaeological Association from un- The third spelaeogriphacean taxon, Acadiocaris derground cave pools in Bat Cave, Table Moun- novascotica, is a Lower Carboniferous form col- tain, South Africa, Gordon (1957, 1960) described lected from the Maritime Provinces of Canadaand

a new malacostracan species, Spelaeo- was described initially by Copeland (1957). It was

griphus This later redescribed Schram who lepidops. crustacean possesses a typi- by (1974), placed it cally peracaridan brood pouch with five pairs of into his newly-erected spelaeogriphacean family

oostegites, and was thus placed within the Super- Acadiocarididae, reassigning it from the peracaridan ouler Peracarida. While its closest affinities seem Order Anthracocaridacea as previously proposed

to be to the , it is not sufficiently simi- by Brooks (1962). This species is described from

lar to any of the existing peracaridan orders for it poorly preserved material, composed of black to be placed comfortably within any of these or- carbonaceous pyritized films on black shale which, ders. Thus, the Order Spelaeogriphacea was erected as Schram (1974) has observed, appears to be in

by Gordon (1957) accommodate this and of to new .the process disintegrating as the pyrite in them

complicated” taxon. Little research has followed oxidizes. This form several appears to posses ‘primi- up on this initial with the tive’ characters when the other description, exception compared to spe- of work on the and respiratory mechanism its laeogriphacean taxa, such as an absence of an optic

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notch in the rectangular carapace, well-developed taxa has also developed on other fronts recently.

and elongate thoracopodal endopods, and five pairs Eva E. Pinardo-Moya at the Unidad de Paleontolo-

ofwell-developed but relatively small and simple gia, Universidad Autonoma de Madrid is currently

the pleopods. working on description of a new spelaeogri-

A fourth species of spelaeogriphacean has now phacean taxon from Lower (Barremian)

been discovered from two equivalent localities deposits of Las Hoyas, Spain (pers. comm.). In

Museum consisting of Upper Jurassic outcrops in Liaoning addition, Gary Poore, of Victoria, Aus- Province (Fig. 1) of the People’s Republic of China tralia and Bill Humphries, Western Australian Mu-

and is the third taxon of a series of new crusta- seum, are describing a newly discovered Recent

cean species from China to be described by the spelaeogriphacean, said to closely resemble

the authors. The first locality represents Jianshangou Potiicoara brasiliensis, from the Pilbara Craton

sedimentary intercalation of the Upper Jurassic of Australia (Humphries, pers. comm.).

Yixian Formation and is found in Jianshangou The phylogeny of this enigmatic order will be

The second discussed in detail later in this village, Chaomidianzi, Beipiao County. some paper. How-

the different will be used than that locality represents slightly younger Dakangpu ever, a approach

sedimentary intercalation, also from the Yixian For- of Pires (1987) who conducted a phylogenetic

mation, and is located in Dakangpu village, Yixian analysis of the orders composing the Superorder

County (Chen, 1988; Chen et ah, 1980). Peracarida. We will use the cladistic method to

Spelaeogriphacean research dealing with new suggest possible relationships within the order. In

I- T'g- of collection sites of (inset: with Locality map Liaoningogriphus quadripartitus, highlighted by pointers China, map area shown as a black rectangle (highlighted by small pointers)).

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will include members no visible of Para- addition, we representative appendages (see descriptions paratypes).

NIGP 126270A/B: lateral view, thorax, abdomen and of the peracaridan orders in this analysis to deter- types, tail fan, with pleopods; NIGP 126275-1: dorsal view, carapace mine the ordinal relationships suggested by our and thorax with antennae and antennules; NIGP 126278A/B- data. 2: lateral view, (incomplete) head, thorax and (incomplete)

abdomen, with partial thoracopods; NIGP 126286-1 and 2:

ventral view, abdomens with incompletetail fans and pleopods;

Systematic palaeontology NIGP 126348: dorsal abdomen and tail fan, partial pleopods; and NIGP 126352: ventral view of complete (juvenile?) ani-

mal. Also figured specimens 126275-8; 126276-8-1,2; 126278A- Class Latreille, 1806 1,3,4. All material is housed at the Nanjing Institute ofGeology Subclass Grobben, 1892 and Palaeontology(abbreviated here asNIGP), Academia Sinica, Peracarida 1904 Superorder Caiman, the People’s Republic of China.

Order Hemicaridea Schram, 1981

Suborder Spelaeogriphacea Gordon, 1957 Additional material.-NIGP 126271: 1-5, NIGP 126272-126274,

NIGP 126275: 2-9, NIGP 126276: 1-7, NIGP 126277, NIGP Family Spelaeogriphidae Gordon, 1957 126278A/B: 5-12, NIGP 126279: 1-20, NIGP 126280-126281,

NIGP 126282: 1-2, NIGP 126283-126285,NIGP 126286: 3-6,

NIGP 126287A/B,NIGP 126349. Liaoningogriphus Shen, Taylor & Schram, gen. nov.

Localities and stratigraphy. - Jianshangousedimentary intercala-

Diagnosis. - with shallow smooth ‘op- Carapace tion, Upper Jurassic Yixian Formation from Jianshangou vil- tical’ grooves and smoothly rounded lateral mar- lage, Chaomidianzi, Beipiao County, Liaoning Province; and

gins extending laterally to third thoracomere, with Dakangpu sedimentary intercalation. Upper Jurassic Yixian second Formation from Dakangpu village, Yixian County, Liaoning thoracomere exposed via a mediodorsal Province (see Fig. 1). indentation; pleomeres 1-5 large, well developed and setose, with large, subequal, subrectangular Description. - The body is elongate and cylindri- protopod, ovoid endopod and ovoid, two-segmented cal and is approximately 3-4 times as long as it is exopod; subtriangular telson with pair of short, wide, with a maximum observed length of >1.75 medial terminal spines; well-developed setose Table PI. The makes cm (see I, Ib). carapace up uropods with rectangular protopod, large, ovoid approximately one fifth of the total body length endopod and exopod of two subequal segments. (see Table I) and is undecorated, thinly sclerotized

and covers the head and most of the first two

- The Etymology. genus name Liaoningogriphus 126271 It thoracomeres (NIGP -1). possesses a very is derived from the name of the region, Liaoning short (<0.5 mm), broadly rounded rostrum. The Province, from which this species has been col- lateral margins are smoothly rounded at lected, carapace and ‘griphos’ (‘something complicated’), both the anterolateral and posterolateral ends of both to reflect the confusing nature of the pleopods the carapace, in an almost ovoid appear- of this resulting and as a dedication to the work of ance in lateral view. Shallow, smooth ‘optical’ Gordon with this group. located between the and grooves are rostrum the

A 0.5 anterolateral margins (PI. Ib, PI. Ild). mm

deep indentation occurs along the medial poste- Liaoningogriphus quadripartitus Shen, Taylor & in Schram rior margin, resulting a small portion of the , sp. nov. (Figs. second thoracomere being exposed dorsally. The 2-3; Pis. l-II)

posterolateral margins partially cover the anterior

of the third (NIGP 126276-8) Diagnosis. - part pereiomere (see As only one species is currently rec- ognized for this Fig. 3). genus, the species diagnosis is the The antennules two robust and same as that for the possess flagella genus. three a peduncle of segments, totalling 2 mm in

Material. The basal is 0.8 NIGP 126269 length. segment approximately Holotype, (PI. lb): carapace, thorax, a domen and almost complete tail fan, preserved in dorsal view, mm long, while the second and third segments are

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Table /. Measurements of spelaeogriphacean morphology, from dorsoventrally oriented specimens (in mm).

Specimen carapace carapace length thorax cephalothorax abdomenwidth abdomen telson width (max) (min/max) width (max) length (max) length (max) length

126269 5.8 2.9/3.4 4.5 7.5 3.8 8.0 2.0

126271 4.9 2.9/3.2 3.8 7.1 3.0 9.0 -

- 126274 -/- - - 3.0 - 2.2

126275-1 - 2.7/3.3 - 7.0 3.1 - -

126276-8-1 4.1 2.7/3.2 3.5 7.0 3.0 - -

126276-11 - -/- - - 3.1 6.1 -

126276-8-2 4.0 2.5/3.5 2.8 5.9 2.5 6.2 1.7

- - 126276-7 - - - 7.3 1.9

- 126275-3 - 3.0 - 3.6 7.6 2.0

126281 - - 3.3 - - - -

126278A-1 - - 3.6 6.8 3.6 7.8 1.9

126275-9 4.4 2.5/2.8 3.2 6.8 "

each approximately 0.6 mm in length. The anten- and 0.75 mm in length (NIGP 126269; NIGP nae are uniramous, with only three segments clearly 126271-1). Specimens preserved in a skewed dorso- visible in and ventrolateral the peduncle. The first second pe- or position show a slight width re- duncular segments are equal in length and together duction in thoracomeres 7 and 8, which probably

the distal in are approximately same length as the reflects the presence ofshorter pleura these seg- segment, with a total peduncular length of2.5 mm ments (Pis. la, lb, lid). These pleura are smoothly

(N1GP 126275-1, 8). No scaphocerite is evident rounded and are directed ventrally (NIGP 126282-

(Fig. 2a). Neither antennal nor antennular flagella 1; NIGP 126271-2).

is known. The from the arc complete, thus flagellar length not following can be determined few

The observed is ~1 far Limbs 2-8 well longest mm long: a cry preserved thoracopods (PI. la). are from those of the recent Spelaeogriphacea, which developed and approximately equal in size, with are almost as long as the total body length. the endopods consisting of 1) a protopod, made

The thorax makes fourth of small 1 in up approximately one up a (approximately mm length com- ofthe Table with of body length of the animal (see I) bined) coxa and basis (details the attachment to the last six free thoracomeres the thorax elon- exposed (Pis. Ib, are, unfortunately, not clear), 2)

Thoracomere size is reduced tho- ischium and and lid). anteriorly: gate subequal (1.0 mm) merus, racomeres 1-3 are smaller than the others and 3) slightly shorter carpus, propodus and dactylus.

in possess medially directed lateral margins, in con- These are subequal length (0.8 mm) but become

of 4-8. trast to the rectangular shape segments In progressively narrower distally. The dactylus is adult individuals, thoracomeres 1, 2 and 3 have pointed distally. Thoracopod 1 is larger than all widths of 3.0 and 3.5 and of others and be into maxil- 2.5, mm lengths 0.5, appears to developed a

0.6 and 0.7 broader mm, respectively (detailed measure- lipede (PI. la), possessing a ischium, merus ments for the anteriormost thoracomeres are and than all other not carpus thoracopods (NIGP

in I due the fact the and provided Table to that these seg- 126278A-1). Both ischium merus are ments can be accurately measured in only two subtriangular. Precise attachment to the thorax is specimens). Thoracomeres 4-8 are subequal in size not determinable, as no clear signs of the coxa or and 4.0-4.5 in measure approximately mm width basis are visible. The propodus is not completely

PI. /. Liaoningogriphus quadripartitus: a), lateral view of thorax with partially preserved thoracopods, poorly preserved head, and

abdomen with incomplete pleopods (paratype; NIGP 126278A-5, *6); b), dorsal view of animal showing carapace, thorax, abdomen and tail fan partial (holotype; NIGP 126269, x5.5); c), ventral view of complete juvenile animal (paratype, NIGP 126352, *24) (al

= first abdominal a6 =sixth abdominal = = = t8 = thoracic segment; segment; c carapace; mxp maxillipede; pi pleopods; eighth

= = te = = segment; telson; tp thoracopod; tp8 eighth thoracopod; u uropod).

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preserved, and no trace of the dactylus is evident. are elongate and biramous and are approximately

No exopods or epipodites are observed on any of twice as long as the telson. Each uropod possesses the thoracopods preserved (PL la, c). a rectangular protopod that is setose along its distal

The abdomen makes almost half of the lateral The setose has arti- up one margin. exopod two total body length (see Table 1). Pleomeres 1-5 are cles, with the rectangular proximal segment equal

in width 3.5 in size and the The equal (approximately mm), with shape to protopod. distal seg- pleomere narrower 6 being slightly narrower(approximately ment is but approximately of same length

3.0 1-4 in mm). Pleomeres are equal length (ap- as the proximal article, somewhat pointed distally, proximately 1.25 mm), while 5 and 6 are slightly and also setose. The endopod is approximately 1.5 longer than the first four (approximately 1.5-1.75 times the size of the uropodal protopod, is ovoid mm) (NIGP 126269; NIGP 126348) (PI. Ilb, c). in shape and heavily setose (NIGP 126278A-4;

Pleura are well developed (PI. Ilb): they are rounded NIGP 126348). and oriented See 3 for dorsal laterally anteriorly but become pro- Fig. complete and lateral recon- gressively more posteriorly directed along the length structions of Liaoningogriphus quadripartitus. of the abdomen (NIGP 126269; NIGP 126278A-

3) (PI. Each of 1-5 - The He). pleomeres possess a pos- Species etymology. species name quadripar- teriorly-pointed triangular apodeme, which is clear- titus (‘four-parted’) is a reference to the two-seg- ly visible but only in dorsally preserved specimens mented nature of the pleopodal exopod, resulting

(PL Hd) suggesting thinly sclerotized tergites (NIGP in pleopods with four ‘articles’.

126271-1; NIGP 126276-8).

Very well-developed, elongate, biramous pleo- Remarks. - As mentioned previously, specimens

pods are of the present on pleomeres 1-5 (PL Ilb, c; Fig. new spelaeogriphacean taxon Liaoningo-

2b). The have been subrectangular protopod possesses a slight griphus quadripartitus collected from mediolateral indentation to the right of the center two separate localities representing the Upper on the otherwise rounded proximal margin. The Jurassic Yixian Formation from the Northern China distal protopod margin is sigmoidally curved, with tectonic platform. The Yixian Basin is a small, iso-

a grooved invagination near the lateral margin, and lated fault basin formed as a result of volcanic anchors a two-segmented exopod (PL 11a) and an activity initiated by the subduction of the Pacific ovoid endopod. The proximal exopodal segment plate under the eastern part of the Asian conti- is subtriangular with a broad distal margin while nent. the distal lobe is The ovoid, creating an overall ovoid exposure at Jianshangou village, Chaomi- appearance for this ramus (Fig. 2b, Pis. 1c, 11a). dianzi, Beipiao County consists of intermediate-

Both the distal lobe and the basaltic volcanic rocks interbedded exopodal endopod are thinly with very quite setose, although setal length is undetermined. fine-grained grey paper shales (Chen et ah, 1982), All three sometimes pleopodal segments are subequal in size slightly calcareous. Associated with the (NIGP 126269; NIGP 126286-1,2; NIGP 126348) spelaeogriphaceans in this strata are the concho- (Fig. 2b). stracan Eosestheria ovata and the fish Peipiaosteus The telson is subtriangular, is slightly longer than panii. This ‘exposure’ is actually the eroded banks wide, possesses a pair ofshort, robust median ter- of a riverbed, and extends several hundred meters minal spines and in Almost three of (NIGP 126348) is subequal vertically. metres this unit are ex- length to the sixth pleomere (PL Ilb). The uropods posed at this locality.

PI. II. Liaoningogriphus ventral view of quadripartitus: a), pleopods (abdomen), arrow suture on the . pointing to exopod (paratype; 126286-1, *9.5); b), dorsal view of abdomen and tail fan with partial preservation of pleopods and pleura of abdominal visible (paratype; NIGP lateral view of and visible with Jergitcs 126348, x9); c), abdomen, pleopods pleura partial tail fan (paratype; 126270B, xg); d), two and incomplete specimens: 1, with complete carapace thorax, incomplete abdomen; 2, with complete * abdomen oiax, and tail two fan, arrows indicate ofthe apodemes mentioned in text (NIGP 126276-8-1,2, x5) (a6 = sixth abdominal

= segment; = pi te = pleopods; telson; u uropod).

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2. and lucida anterior end of Fig. Liaoningogriphus quadripartitus: a) photograph corresponding camera drawing showing carapace

and = = = details ofantennae and oneantennule; a antennule, an antenna, c carapace margin (NIGP 126275-1, *14); b), photograph and view of of associated = corresponding camera lucida drawing of ventral single pleopod plus portions closely pleopods; p

= protopod, x exopod, n = endopod (NIGP 126286-2, *23).

The second locality represents the slightly stones (Chen, 1988; Chen et ah, 1980) and slightly younger Dakangpu sedimentary intercalation and darker silty limestones, and is an intercalation of is found in Dakangpu village, Yixian County (Fig. intermediate-basaltic volcanic rocks (Chen et al., is 1). It composed of thin-bedded fine-grained silt- 1982). The insect Ephemeropsis trisetalis and the

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the calities are located near southern margin of what is interpreted as humid temperate zone bor-

dering on a semiarid-arid subtropical zone (Wang,

1985).

Preservation varies considerably betweenthe two

localities. Specimens from the siltstone locality at

Dakangpu village are preserved as carbonaceous

films with reliefevident in no any specimens. Some

in details of internal anatomy are observable dorso-

oriented ventrally specimens, with both upper and

lower preservational surfaces at least partially

visible in most of these specimens.

Almost all of those collected from the paper

shales at Chaomidianzi show some relief. Many

of these Chaomidianzi specimens are preserved

as molds, with no traces of the original organic

material (or replacement material) evident. Oth-

ers are partially phosphatized; these show consid-

information erably more than the matrix-impres-

sion specimens, usually possessing greater relief

as well. No internal anatomy is visible; however,

in some specimens the outlines of structures (i.e.,

segment margins) on the lower preservational

surface visible the are through upperpreservational face of the specimens.

Fig- 3. Lateral (A) and dorsal (B) reconstructions ofLiaoningo- There is (and has been for some time) discus- griphus bar = 0.1 quadripartitus. Scale cm. sion and debate to the as actual age of the Yixian

Formation. The traditional perspective, as used in conchostracan genus Diestheria are also found at this paper, is that it was deposited during the Upper this each in abundant locality, amazingly numbers Jurassic. However, recent age dating studies (i.e.,

(Zhang et of 1995 40 39 al., 1976). Three specimens previ- the Smith et al., study using Ar- Ar) for

ously undescribed Yixian terrestrial insects (tentatively the Formation suggest that it is actually identified thus far as a cicada, a necopteran and a Early Cretaceous in age. An attempt will later be

havealso hymenopteran) been found at this local- made to help clarify this issue through an exami-

ity, which will be the focus of later This of the a paper. nation palynomorphs present in material exposure is 10 small, a lens of approximately cm collected from the Yixian Formation. This infor-

thickness that is in exposed along the banks of a mation will be included the previously men- railroad track excavation. tioned paper dealing with several new insect taxa This region contains a freshwater fauna known from the region. as the Jehol Fauna, which includes the previously mentioned insects, conchostracans and fish and extends the through Late Jurassic to the early Creta- Phylogenetic analysis ceous in eastern Asia. Recent exploration through- out the area has in- uncovered numerous new taxa A phylogenetic analysis was undertaken to estab-

cluding dinosaurs, birds and angiosperms. The dish the relationship of the new Chinese spelaeo- stratigraphy and of this lithology region, as well griphacean genus/species to the other known forms. as the associated tell faunas, us these were The show derived deposits two recent species many char- aid down in a lacustrine environment. These lo- acters not present in the plesiomorphic Carbon-

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Table II. Data matrix used in the phylogenetic analysis discussed sis and concluded that the Thennosbaenacea and

in this Character for information paper (see descriptions regarding Mictacea formed a sister clade to the Spelaeogri- the identity ofthe characters listed here, indicates inapplicable phacea, and that these taxa together formed a sis- character, “?” indicates uncertain or unknown datum). the ter group to , Tanaidacea, Taxa Characters and . Thus, at one time or another,

almost has been every peracaridan taxon suggested 00000000011111111112222 by cladistic analyses to be the sister taxon for the 12345678901234567890123 Spelaeogriphacea.

Although the sister of the Spelaeogriphacea Mictocaris halope 0-000001300100100002100 group

is not there seems to be Halosbaena acanthura 0-200211102220000010000 evidently clear, some agree-

Acadiocaris novascotica 0-2012112000002200202070-20121120000022002020? ment as to the “cluster” of taxa most closely re-

SpeheogriphusSpelaeogriphuslepidops 11211111211100211021101 lated to the Spelaeogriphacea. Thus, for our anal-

Potiicoara brasiliensis 10111011200001231022111 ysis we included relatively plesiomorphic mem-

Liaoningogriphus quadripartitus 0-10027121001012001111? bers of each of these four peracaridan taxa to act NeotanaisNeotanais micromorpher 10211021101222000024-12-001101222000024-12-001 as representatives of the larger group: the mictacean, LeuconLeucon bacescui 1022002311222024-100001 Mictocaris halope; the thermosbaenacean, Halos-

baena acanthura; the tanaid, Neotanais micromor-

pher; and the cumacean, Leucon bacescui. iferous species Acadiocaris novascotica, and thus the affinities of the time-wise intermediate form

from the Jurassic are of interest. A matrix of 23 Character descriptions characters (see Table II), based exclusively on

was coded Mac- morphological characters, (using In developing our character matrix, we selected

Clade 3.01) for an analysis using PAUP 3.1.1. features that would minimize the effect of Spec- ques- imens of and Potiicoara Spelaeogriphus lepidops tion marks for unknown, or dashes for inappro- brasiliensis with the of Antonio (obtained help priate, scorings. We choose to emphasize, but not

Carlos and Norma Sharratt) as well as those features that could be Marques exclusively so, scored the literature data for the other for this previous provided all taxa. We feel is justified, however, on The matrix spclaeogriphacean taxa. resulting pos- the basis of the need to incorporate information sessed almost no missing data, a relative rarity in from fossils into The any phylogenetic analysis. cladistic fossil taxa. analyses utilizing characters we used, with some explanations con-

The sister to the group Spelaeogriphacea remains cerning them, are provided here. a vexing issue. Pires (1987), through an ordinal

level cladistic of the Peracarida, concluded 1. has analysis Ocular lobe. This feature a relatively lim-

the that the Mictacea are most likely sister ited within the of the group. application context groups Other authors, however, have determined differ- we are analyzing. Extension beyond these taxa ent for the sister-group relationships Spelaeogri- would have required us to deal with another char- The of Schram phacea. analysis (1981) suggested acter expression not encountered within the taxa the order Thennosbaenacea as the sister to group we focus on in this study, viz., stalked compound a clade the orders Cu- including Spelaeogriphacea, eyes. We are assuming in this case that ocular lobes, macea and Tanaidacea(Order Hemicaridea Schram, even without optic elements, are in fact homologs

1981), a similar conclusion to that drawn Wat- of stalked by eyes. Presence of the lobes themselves ling (1981, 1983). Schram, however, concluded can be difficult to interpret. In certain ingolfiellid that the and Tanaidacea are sis- Spelaeogriphacea amphipods, the lobes are so small as to be almost ter whereas the Cumacea groups, Watling suggested undetectable. This condition is especially relevant and Tanaidaceaare the sister to the groups Spelaeo- because apparent lack of such lobes in the fossils griphacea. Wagner (1994) undertookhis own analy- could be due to vagaries of preservation. We pre-

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in fer this case record whether or not exhibit simply to not we any great development of the groove.

can see detectable lobes. (Absent = 0, present (Absent = 0; present =1.)

= 1.)

6. Antennalscale. The variable development of the

2. The feature deals with Ocular lobe margin. some antennal scale is often employed in phylogenetic

variations in of the we can note the decoration lobe analyses of eumalacostracans. Its use is problem- itself. The exact function of such papillations for atic. The functional meaning of the feature remains

now remains uncertain. This feature remains in- unclear, although in eucarids and mysidaceans it

formative, since it this time role however, appears at may serve some as a stabilizer in swimming.

to be an autapomorphy of Spelaeogriphus lepidops. The antennal scale is absent in all mictaceans, thus

(Distal margin smooth = 0; distal margin papil- it seems to associate thermosbaenaceanswith the

lated = 1.) hemicarideans. Note, however, that neither Potii-

the the tanaidacean coara, cumacean, nor in our 3. Rostrum. Use of the rostrum be con- matrix such scale. is term might possess a It a highly variable

sidered too a term in this Cer- condition the chose “strong” context. among Tanaidacea, e.g., we

in of these tainly nothing any taxa approaches the Neotanais micromorpher as our tanaidacean, which

large, rostra noted in well-developed decapod lacks a scale, but a tanaid-like Apseudes hermaphro-

We = eucarids, mysidaceans, or euphausiaceans. diticus possesses one. (Absent 0; small, i.e.,

the term refer of employ rostrum to to any antero- shorter than, length first peduncle segment of

median of the shield. All = development carapace antenna 1; large, i.e., equal to or longer than the taxa in this = analysis possess some development length of first peduncle segment of antenna 2.)

in this = = area. (Absent 0; broad, rounded 1; short,

pointed = 7. This 2.) Antennalpeduncle. feature is a synapomor-

phy of the and Hemicaridea. (3 4 -Anterior The = = branchiostegal development. segments 0; 4 segments 1.)

of branchiostegites the carapace can form promi-

nent features the eumalacostracan and 8. Pereiomeres with the amongst fused carapace. Degree

especially of of the peracarid . They appear to development carapace over the thorax be linked of with specializations the underlying and its fusion to the underlying segments is an limbs for respiration. The hemicarideans use the important architectural feature of peracarid body space beneath as a respiratory chamber ventilated plans. All of these share this feature to

by actions of the large bailer on the maxillipedes. some degree, i.e., at least the first maxillipedal

This character is fused. The functions as an apomorphy for segment so degree of further fusion

in and hemicarideans, although our Chinese species does seen cumaceans tanaidaceans is linked with

not to other appear exhibit any great development of specialization of anterior thoracopods in this = region. (None = = evident 0; present 1.) addition to the first as maxillipedes. (None 0;

one = 1; two = 2; three = 3.) 5. Cervical groove. This is a common feature on

eumalacostracan 9. Number carapaces, marking the point at of thoracomeres covered by the cara- which the shield fused the The number of thoracomeres carapace to underlying pace. covered by the cephalic into segments passes the unfused portion carapace as opposed to the number fused to the overlying the thoracic segments. The cervical is cephalon are separate issues. In the larger context

the most common of the different carapace grooves, of a eumalacostracan, or even malacostracan analy-

although eucarids can whole this display a complex sis, feature takes on more importance. Its high and diagnostic of such furrows. array This feature degree of variability here, however, probably does delineates hemicarideans, the Chinese to although not effect the analysis other than help serve to form and the cumacean used in and this analysis do unite hemicarideans suggesting a shared fea-

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ture with tanaidaceans. (Eight = 0; three = 1; two clade. (Margin entirely setose = 0; distolateral +

= 2; one = 3.) distal setae/spines only =1; distal setae/spines only

= 2; distolateral setae/spines only = 3.)

10. Thorax/abdomenproportions. A variable fea-

it effect here in size. This character has ture, has perhaps limited ordering 17. Terminal telson spine

taxa. It is retained, however, for possible use in some potential for distinguishing features within

the The usefulness distinguishing among the spelaeogriphaceans. spelaeogriphaceans. can only

(Thorax same lenghth as abdomen = 0; thorax < be assessed, however, as more taxa are discov-

abdomen = 1; thorax > abdomen = 2.) ered within this order. (Prominent = 0; short and/

or thin = 1.)

11. Pleopod 5 rami. The next few characters are

problematic. They entail major architectural fea- 18. Pleotelson. This feature is a synapomorphy

tures of the body plan but often exhibit wide ranging uniting the non-spelaeogriphacean hemicarideans.

variation between taxa. They are nevertheless (Absent = 0; present = 1.)

employed fairly consistently in the literature. In

this the variation in the characters 19. This is a feature that seems particular case, Uropodal endopod.

noted for the in be effective in a between are, most part, autapomorphic ex- to defining relationship

pression. We retain this feature here since its rel- thermosbaenaceans and hemicarideans, and hold

for within evance in the context of some future analysis us- promise sorting relationships the latter.

discovered fossil = than first ing newly or recent taxa may (0-2-Segmented 0; longer exopodal

become more evident. (Biramous = 0; uniramous segment = 1; equal to first exopodal segment = 2.)

= 1; absent = 2.)

20. Uropodal exopodal segments. This is a vari-

12. Pleopod 5 development. The degree of devel- able feature shared between mictaceans and

of is which be useful opment a limb, we believe, largely independent spelaeogriphaceans, may greatly

from the variety of form as expressed in character in future analyses involving Spelaeogriphacea.

I I. The same comments that pertain to character (Proximal segment longer than distal = 0; subequal

II apply here as well. (Well developed = 0; re- in size = 1; distal segment longer than proximal =

duced = 1; absent = 2.) 2.)

13. Pleopod exopod. The potential significance of 21. Uropodal protopod. Again, the subtle differ-

a distinctive character such as this will be only ences in proportions of uropod and telson elements

hold in fully realized as more taxa are added to the ma- may promise establishing relationships

trix. (A single article = 0; 2 articles = 1; exopod i between spelaeogriphaceans. They can be easily

absent = 2.) evaluated on fossils as well as recent forms. (Longer

than first exopodal segment = 0; equal to first

= 14. Telson terminus. At present this character is exopodal segment 1; shorter than first exopodal

an autapomorphy for Potiicoara brasiliensis. (Sim- segment = 2.)

ple = 0; decorated = 1.)

22. General form of uropod. This feature now

15. Telson size. This is a fairly consistent feature. functions with some potential for sorting relation-

It finds some utility in clarifying relationships within ships within spelaeogriphaceans. (Length > width

the = ~ = hcmicarideans. [Long (equal to exopod at the 0; length width I.)

= diaeresis) 0; medium = 1; short (equal to length of exopodal protopod) = 2.] 23. First maxillipedal epipodal bailer. This fea-

is first ture added here as a step to resolving rela-

16. Telson decoration. the within the hemicarideans. It would marginal Again utility tionships ap- of this feature functions within the hemicaridean pear to act as a synapomorphy shared by mem-

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bers remains difficult assess of the group, but it to for fossil members. It may be capable of finer definition, however. Cumaceans seem to have digitiform bailers, and tanaidaceans generally have their bailers developed as large membranous lobes.

The epipodite in Spelaeogriphus is rather cup-like,

Our but that in Potiicoara is described as “oval”.

SEM study (see Discussion) hopes to resolve this feature further. (Not greatly modified= 0; present

= 1.)

Results

The outcome of this analysis is presented in Fig.

4: 54 a single tree with a length of steps and a

relatively high Consistency Index (Cl) of 0.74. The

four in spelacogriphacean taxa occurred as taxa the same part of the tree, suggesting the presence of some phylogenetic affinity between these taxa.

Interestingly, L. quadripartitus surprisingly ends

up as the basal-most of the four spelaeogriphacean

taxa. When these four taxa are analyzed alone,

however, A. ends as the basal-most novascotica up

taxon = 24 Cl = in the tree (tree length steps, 0.88; Fig- These 5). somewhat contradictory results sug-

gest that L. hand quadripartitus possesses on one

some features that unite it with the recent Spclaeo- griphacea (more so than A. novascoticd), but on the other some that are more typical of the non-

spelaeogriphacean peracarids. Thus, it takes up an

ambiguous ‘intermediate position’ in the overall

peracaridan tree.

Perhaps even more notable, however, and cer-

of tainly more far-reaching phylogenetic import, is the result that nested within this “particular hemi- caridean clade” are the Tanaidaceaand the Cuma-

cea as sister This confirms earlier groups. the re- sults of Watling (1983) and further suggests that the Spelaeogriphacea as it currently stands may actually represent a paraphyletic group. It is important to stress here that the intention

°f this analysis is not to provide a definitive pic- ture for the phylogenetic relationships between the peracaridan taxa. It is intended to demonstrate that

phylogenetics can be a useful tool even at the al-

pha taxonomic level, as it may suggest relation- Fig. 4. The single tree obtained from a phylogenetic analysis ships that are not visible from always - immediately of the data matrix shown in Table II (tree length 54 steps, Cl observations of morphology focused as they often - 0.74).

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either tion relevant to possibly establishing a dis-

tinct spelaeogriphacean clade, or confirming the

the paraphyletic status of Spelaeogriphacea sug-

gested herein. This kind of research, as well as

Re- the anticipated description of new fossil and

cent taxa, will go a great distance in helping to

resolve the taxonomy of these and other closely related peracaridan orders.

Unfortunately, several obstacles continue to stand

in the way of the determination of a definitive

for this of the avail- phylogeny group. First, some

able information is not particularly informative for

an analysis of the Peracarida. As can be seen in

the data matrix used for our analysis here (Table

II), or by examining representative taxa from each

of the respective orders and suborders discussed

here (Schram, 1986), many of the “obvious” mor-

phological characters used to define these taxa are

shared with most if not all of the other peracaridan

orders. For example, most members of these five

orders bodies with possess elongate, sub-cylindrical

reduced that covers the anterior- a carapace only

most region of the thorax. Is this character “facies”

merely plesiomorphic, or is it a convergent set of

features that in connection with appears a particu-

lar set of habitats? At this point we do not know. Fig. 5. The single tree obtained from a phylogenetic analysis of the characteristic features of Second, many ofonly the spelaeogriphaceantaxa in the matrix shown in Table each we can employ actually at this II (tree length - 24 steps, Cl - 0.88), species appear

time to be autapomorphies and thus uninforma-

in lens that entails tive a phylogenetic analysis, the bi-lobed are through a some preconcep- e.g., ofL. tions about taxonomic status. pleopodal exopod quadripartitus, orthepapil-

lated distal margin on the ocular lobe ofS. lepidops.

It is conceivable, however, that as more spelaeogri-

of these features Discussion phaceans are discovered, some

be shared with these be found may yet to taxa.

their inclusion here for the time Ongoing phylogenetic research Nevertheless, being remains problematic.

We Third, several features used in recently obtained some limited material for commonly pre- vious discussions of both Potiicoara brasiliensis and Spelaeogriphus peracaridan phylogenetic re-

well the often cannot be determined from fos- lepidops, as as specimens of soon-to-be- lationships

information describcd species from Australia, with the inten- sil material, e.g., optic anatomy, or tions of blood and conducting the first SEM based compara- concerning excretory systems (Watling,

1983). To what extent this kind of information future tive analyses of these species. Such studies should

the analyses of phylogeny can incorporate without reveal further details about anatomy of these

that will lead compromising any phylogenetic signal with high species hopefully to greater taxo- for levels of remains unclear. nomic refinement these taxa with some added uncertainty boost Finally, of the known peracaridan taxa for understanding their phylogenetic rela- many in This live in habitats that are not often the tionships. study may uncover some informa- preserved

Downloaded from Brill.com10/11/2021 09:56:06AM via free access Contributions to Zoology, 68 (1) - 1998 33

fossil record, such as the ground-water habitats zil, S. lepidops in South Africa, and the as-yet- discussed here, and the deep-water habitats in which unnamed spelaeogriphacean from Australia. How-

taxa the each of the known fossil taxa is in many peracaridan live, e.g., bathynella- ever, located

baicalensis and B. which the northern with A. found ceans Bathynella magna, hemisphere, novascotica live in Lake Baikal at depths of up to 1440 m, or in eastern Canada, L. quadripartitus in China, and the many hemicarideans known from the deep sea the as-yet-unnamed Cretaceous form in Spain (Fig.

As with (e.g., see Schram, 1986). a result, the informa- 6). Schram (1977, 1982), only one fossil

from the in tion available fossil record will always species the Carboniferous, suggested an early be incomplete, resulting in ambiguities in any data Laurentian origin with later Gondwanian ‘diver- matrix constructed. sification.’ This pattern of Laurentian origin with

Due to factors such as these, problems will Gondwanian dispersal (most likely occurring at continue occur with resolve the the time of the formation of the to any attempt to Pangean phylogenetic relationships among all peracaridan supercontinent in the ) seems reasonable taxa. Whether these problems will remain insur- and finds reflection perhaps in other malacostracan mountable remains to be seen. taxa. The new species from Australia would seem

to support the prediction of Schram (1974) that

with further research, living spelaeogriphaceans

Biogeography would be found “in the areas.”

Nevertheless, the new Cretaceous form from

The number of of is in combination with our Late Jurassic species Spelaeogriphacea Spain, spe-

their distribu- cies from an alternative unfortunately sparse. Despite this, China, might suggest sce- tion suggests a potentially interesting set of bio- nario for the evolution of Spelaeogriphacea. Might

the geographical scenarios. The known recent forms we look to bathynellacean syncarids for an-

are restricted to the southern hemisphere, suggesting other model? The distributional pattern of all

their is of Gond- with their late geographic pattern a remnant a spelaeogriphaceans relatively per-

wanian distribution: P. brasiliensis found in Bra- sistence in northern continents, could be one of

I ig. 6. World map showing distributions of Recent (indicated by ‘o’) and fossil (indicated by ‘x’) spelaeogriphacean taxa.

Downloaded from Brill.com10/11/2021 09:56:06AM via free access 34 Shen Yan-bin, R.S. Taylor & F.R. Schram - Fossil speleaogriphacean (Crustacea) from China

world-wide ubiquity. Under such a scenario we Onderzoek’, Nederlandse Organisatie voor Wetenschappelijk

Onderzoek (the Foundationfor Geological, Oceanographic and would be tempted to predict that future discover- Atmospheric Research, the Dutch Organization for Scientific ies of spelaeogriphaceans will not be confined to Research). This project has been partly supported by the Chinese Gondwanalocalities, but will conform to a distri- Academy of Sciences as well (grant no. KZ951-B1-410-2).

bution more akin to that shown the Thanks also extended reviewer and Dr. by bathynella- are to an anonymous to

comments. ceans. These crustaceans are currently found in G.F. Wilson for their valuable

ground-water habitats world-wide, and are believed

to have originated in the Laurentian from a primi-

tive syncarid eumalacostracan in the late Palaeozoic. References

The bathynellaceans later dispersed throughout

Brooks HK. 1962. The Palaeozoic Eumalacostraca ofNorth northern and southern regions with the formation America. Bull. Amer. Paleontol. 44 (202): 163-338. of Pangaea in the Permian (Schram, 1986). We Chen Peiji 1988. Distribution and migration of Jehol fauna believe that such distribution a pattern may even- with reference to non-marine Jurassic-Cretaceous bound-

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