2.6 Reproductive System

Edith Cowan University

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EVOLUTION,SYSTEMATICS & GEOGRAPHIC PARTHENOGENESISOF Ilyodromus (CRUSTACEA,OSTRACODA)

Rylan James Shearn, BSc Hons

A thesis submitted to Edith Cowan University in accordance with requirements for the degree of Doctor of Philosophy

Submitted 27th February 2015 A NRDCINT MORPHOLOGY TO INTRODUCTION N (C RUSTACEA FTHE OF O , C YPRIDOIDEA STRACODA

C HAPTER 2 ) 2.1 Abstract

Several chapters of this thesis frequently use specialist terminology to identify and describe morphological features of ostracods from the superfamily Cypridoidea. To assist the non-specialist reader, this chapter guides the reader through a series of illustrations that serve to explain the fundamental terms and concepts required to interpret these descriptive sections. It is hoped that this chapter will serve as a basic guide, or atlas from which readers can refer back to while reading the remainder of the thesis.

36 CHAPTER 2. THE CYPRIDOIDEA 2.2 Introduction

Ostracods are small shrimp-like crustaceans enclosed by a calcitic bivalved carapace, essentially resembling a mussel with a shrimp inside (De Deckker, 1995). They have been discovered in nearly all aquatic environments (marine and non-marine), deep subterranean waters (Karanovic, 2007), and semiterrestrial habitats (moist leaf litter; Pinto et al., 2003, 2004), while some are commensal on other aquatic organisms (Horne et al., 2002). Many species are generalists, having mainly detrital and herbivorous feeding habits (Martens, 2001), but some predate other small crustaceans (Martens, 2001) or ﬁlter feed (Karanovic, 2012). They are also known to serve as food for ﬁsh (Kornicker and Sohn, 1971; Vinyard, 1979) and waterbirds (Green et al., 2008). Most marine ostracods reproduce sexually (Cohen and Morin 1990), while in non-marine Ostracoda, there are three known reproductive modes accord- ing to Martens et al. (1998):

1. Sexual; whereby females of a species require insemination by a male in order to reproduce, this life cycle involving an alternation between meiosis (producing haploid male and female gametes) and syngamy (fusion of the two gametes to restore a diploid zygote).

2. Parthenogenetic (here and throughout this thesis also referred to as asexual, or asexuality); whereby females produce offspring without male fertilisation or syngamy.

3. Mixed reproduction; whereby populations can consist of sexual, parthenogenetic, or a mixture of both modes (sexual males, sexual females, and parthenogenetic females).

Usually, parthenogenetic females and sexual females of the same species are morphologically indistinguishable. However, sex ratios can be indicative of reproductive mode, whereby any ratio close to 50:50 is likely to be sexual, while populations with a mixture of both modes are likely to have female- skewed ratios (Martens et al., 1998). Females lay eggs on plants, at the sediment surface, or in some species are kept in brood cavities within the carapace until juveniles become fully independent (Martens 2001). For many species, eggs are resistant to desiccation, and can be viable for over 50 years (Martens, 1994a,b), enabling them to persist in highly temporal habitats (See Chapter 6). This also enables ex- cellent dispersal ability for some species, as desiccation resistant eggs can be swept long distances by the wind (Vanschoenwinkel et al., 2008a), go against

2.2. INTRODUCTION 37 wind direction by surviving a passage through the gut of travelling waterbirds (Green et al., 2008), or attaching externally to travelling birds (De Deckker, 1977; Horne and Smith, 2004) or mud wallowing animals (Vanschoenwinkel et al., 2008b, 2011). There are three main superfamilies of non-marine Ostracoda (Cytheroidea, Darwinuloidea and Cypridoidea). Cypridoidea represents over 75 % of the approximately 2000 known extant species of non-marine Ostracoda (Martens et al., 2008), including species of Ilyodromus. Many taxa within the superfamily Cypridoidea, including Ilyodomus, are discussed and described throughout this thesis using specialised terminology. To assist non-specialist readers, this chapter aims to briefly explain basic terminology used in describing extant cypridoidean ostracods. As most of the information given in this chapter is attributed to only a small number of key texts, a different referencing style has been adopted to avoid repetition and to give an overall improvement of readability. Through- out this chapter, unless specific references are given, explanations of terminology were derived from the following sources; Meisch (2000) and Van Morkhoven (1962) for valve structure, Broodbakker and Danielopol (1982), Karanovic (2012), Martens (1987), and Meisch (2000) for appendage structure and layout, and McGregor and Kesling (1969a,b) for reproductive systems. This guide is largely intended to be a visual experience, and the reader is encouraged to interpret morphological concepts mainly through examina- tion of the figures listed at the end, under the guidance of the main text. Some illustrations were made specifically for this chapter, others were adapted from existing illustrations in other chapters or other literature, and have been refer- enced accordingly. Readers can refer to several other useful texts for comparative morphology among the other superfamilies of non-marine Ostracoda in Karanovic (2012) and Meisch (2000), and among other taxa of marine Ostra- coda in Horne et al. (2002).

2.3 Orientation

Like all ostracods, cypridoideans are small bivalved crustaceans, being completely enclosed between two calcite valves. For orientation, these animals are best examined in lateral view, and often an eye (ocular lens) can be observed beneath the carapace in the antero-dorsal region (Figures 2.1 and 2.4), otherwise an antennule may be seen protruding between the two valves, also in the antero-dorsal region. Features toward the core of the animal are referred to as ‘medial’ or ‘proximal’ while features toward the extremities

38 CHAPTER 2. THE CYPRIDOIDEA are usually termed ‘distal’ (Figure 2.1). Carapace length, height and width measurements are usually taken as the maximum distance between parallel points at the anterior and posterior, dorsal and ventral, and right lateral and left lateral margins respectively (Figure 2.1).

2.4 Carapace and valves

During dissection, the left and right valves that constitute the carapace are separated and the inner structures can be observed, which are important taxonomic features (Figure 2.2). Both valves have an inner and outer lamella, that are separated by the vestibulum (Figure 2.3). Some of the inner lamella is calcified around the margin, and the boundary between this calcified and non-calcified inner lamella is well defined and termed the inner margin (Fig- ures 2.2 and 2.3). The calcified part can have inner lists and selvages (Fig- ures 2.2 and 2.3) that have a gasket-like function, often completely sealing the carapace when the valves are closed together, and the variation of these structures are also important taxonomically. The outer margin refers to the outer most edge of the valve in lateral view. Toward the valve outer margin, the outer and inner lamellae may be fused, and here, radial pore canals can often be observed (Figures 2.2 and 2.3). The outer lamella also has normal pore canals that protrude setae on the outer side of the valve (Figure 2.3). The left and right valves are attached to one another by a dorsal hinge and the ostracod closes the valves together with centrally and dorsally positioned adductor muscles (Figure 2.4). These muscles leave attachment scars on the valves (Figure 2.2) called central muscle scars, and dorsal muscle scars, both of which are important taxonomically.

2.5 Appendage layout and terminology

Cypridoidean ostracods have eight sets of paired appendages (Table 2.1 and Figure 2.4) not including copulatory organs (the male hemipenis or female genital lobe). From anterior to posterior, they are the Antennule (A1), Antenna (A2), Mandibula (Md), Maxillula (Mx), Fifth limb (L5), Sixth limb (L6), Seventh limb (L7) and Caudal rami or ramus (CR). Although these names are generally accepted in current literature, other terms are used for the same appendages of cypridoidean ostracods, or for homologous appendages in other ostracod taxa (Table 2.1). The Antennule is the most anterior appendage and is mainly used for lo- comotion, bearing many natatory (swimming) setae (Figure 2.5), but it also

2.4. CARAPACE AND VALVES 39 Table 2.1: Terminology used throughout this chapter and the thesis to refer to appendages of species of Cypridoidea, listed from anterior to posterior, also showing abbreviations used throughout the thesis, and other synonymous terminology used either for different superfamilies of Ostracoda, or by some authors for Cypridoidea. After Horne et al. (2002)

TERMINOLOGY ABBREVIATION OTHERTERMS ADOPTEDHERE Antennule A1 antennula ﬁrst antenna

Antenna A2 second antenna

Mandibula Md mandible

Maxillula Mx maxillule maxilla ﬁrst maxilla

Fifth limb L5 ﬁrst thoracic leg maxilliped maxilla second maxilla walking leg

Sixth limb L6 second thoracic leg ﬁrst thoracic leg walking leg

Seventh limb L7 third thoracic leg second thoracic leg walking leg cleaning leg

Caudal rami CR rami furca furcal rami uropods

has a sensory function, bearing aesthetascs (sensory organs). The appendage has eight segments in most described species of this superfamily (Figure 2.5), with the ﬁrst two often being fused and counted together (reviewed in Kara- novic, 2012). However, many subterranean species have a reduced number of segments. All segments are numbered for explanatory purposes in Figure 2.5, but in subsequent chapters of this thesis, the ﬁrst two are considered a single segment, making a total of seven. The lengths and widths of setae and

40 CHAPTER 2. THE CYPRIDOIDEA claws on the Antennule, although often not illustrated, can be useful taxonomic characters (See Chapters 3 to 6). The Antenna (Figure 2.5) is positioned behind the Antennule (Figure 2.4) and can also have a swimming function, bearing some long natatory (swimming) setae. However, some benthic species have very reduced setae, and this appendage assumes more of a burrowing or walking function (See Chap- ter 5). This appendage also has some sensory function as it bears aesthetascs (sensory organs). Structurally, the Antenna consists of two fused protopodal1 segments, a small exopod2, and a three segmented endopod3 (Figure 2.5). There are usually four setae on the inner side of the second endopodal segment called t-setae (T1-4). More distally, there are three setae (z1-z3), usually three claws (G1-G3) and a small aesthetasc (y2). These structures are sexually dimorphic to varying degrees, and in males, the z-setae can be claw-like, while the claws G1-G3 can be reduced (See Martens, 1987). The third endopodal segment is terminal, short, and has up to two claws (GM and Gm), Gm sometimes being reduced and seta-like. The third endopodal segment also has up to three setae, one of which is a sensory aesthetasc (y3) which can be fused at the base with another seta (Figure 2.5). The Mandibula (Figure 2.6) is positioned behind the Antenna, and is com- prised of a robust coxa that bears teeth that help to grind food, a palp with many setae that have a feeding function, and a branchial plate for respiration. The palp is composed of a single protopodal segment and three endopodal segments. The endopod carries many setae and claws that vary in structure and length, of particular importance are the α, β, γ, S1 and S2 setae (Figure 2.6). The Maxillula (Figure 2.7) is positioned behind the Mandibula (Figure 2.4) and consists of a two segmented endopod (or palp) and three endites4 that function in feeding, and a large exopod that takes the form of a branchial plate for respiration. Although the number of rays on the respiratory plate (Figure 2.7) can be an important character for higher taxonomic levels (Smith et al., 2005), they are often damaged during dissection and thus should be used with caution in taxonomy (reviewed in Karanovic, 2012). The Fifth limb (Figure 2.8) is positioned behind and slightly below the maxillula (Figure 2.4). This appendage is made up of a protopod, an exopod, and an endopod. The protopod bears one or two setae close to the base

1PROTOPODAL (PROTOPOD): The basal part of a crustacean biramous (two branched) limb 2EXOPOD: The outer branch of a crustacean biramous (two branched) limb 3ENDOPOD: The inner branch of a crustacean biramous (two branched) limb 4ENDITES: Additional lobes on the inner margin of the protopod of a crustacean biramous (two branched) limb

2.5. APPENDAGE LAYOUT AND TERMINOLOGY 41 of the appendages called the a-setae, another three setae can be found more distally that are called b, c and d-setae, but can also be missing in some groups (for example the c-seta is missing in all Ilyodromus Figure 2.8). The protopod terminates with a number of setae that are used for feeding. The exopod is a small branchial plate that points posteriorly, functions in respiration, and has up to six rays (Figure 2.8). The endopod has strong sexual dimorphism; in females it is often more simple, pointed and terminating with setae, while in males it assumes the form of a clasping organ (Figure 2.8) that is used to manipulate female valves during copulation. The Sixth limb (Figure 2.9) is a walking leg that is based behind (more posteriorly to) the fifth limb (Figure 2.4). The appendage consists of a protopod that is incompletely divided into two segments, and a four segmented endopod (Figure 2.9). Each of the protopodal segments can bear a seta anteriorly (called d1 and d2) as can the first two endopodal segments (called seta e and seta f), while the third endopodal segment can have two setae (called g setae). Sometimes segments two and three can be fused. The terminal (fourth endopodal) segment is shorter and bears one long claw (called h2) an anterior seta (called h1) and another seta posteriorly (called h3) which can sometimes be more developed and claw-like (See Ilyodromus dikrus in Chapter 4). The lengths of claws, setae and segments can vary between taxa and thus are useful taxonomic characters. The Seventh limb (Figure 2.9) is based behind the sixth limb, but usually pointed dorsally (Figure 2.4), and is used by the cypridoidean ostracod for cleaning the space within the posterior of the carapace. The appendage has a single protopodal segment (Figure 2.9) that can bear two setae anteriorly (called d1 and d2) and another posteriorly (called dp). The endopod has up to two four segments, the first is usually the most elongated and can bear a seta anteriorly (called the e-seta). The second and third endopodal segments are often fused, and can have an anterior seta (called the f-seta) at their fused boundary, and another at the end of the third segment (called the g-seta; missing in illustrated species). Often the terminal (fourth endopodal) segment is also fused to the third segment and forms a pincer organ (Fig- ure 2.9), or may be clearly defined as another segment (reviewed in Karanovic, 2012), but in either case bears three setae of varying structure and length (Fig- ure 2.9). The Caudal rami (singular: ‘Ramus’) are two elongated rod-like appendages that are based at the very posterior of the body (Figure 2.4). The swing together along an arc in a single dimension below the animal, mainly as a walking action, but this action can be vastly accellerated in at least one

42 CHAPTER 2. THE CYPRIDOIDEA genus, enabling a ‘springtail’ escape jump (Matzke-Karasz et al., 2014a). The Caudal rami are not segmented (Figure 2.10), and bears claws and setae at the distal end for many groups within Cypridoidea. These are named with the ab- breviated terms Sa (anterior seta), Ga (anterior claw), Gp (posterior claw) and Sp (posterior seta). However, the morphology of these structures is highly variable, and some setae can assume the form of a claw (as in Figure 2.10). In some groups of Cypridoidea, the rami can be missing, or very reduced (reviewed in Karanovic, 2012). This high variability makes the Caudal rami (or ramus) useful in taxonomy. Additionally, part of the exoskeleton connected to the Caudal rami is called the Caudal rami (or ramus) attachment (Figure 2.10), and its morphology is often used to differentiate between higher taxonomic ranks.

2.6 Reproductive system

In comparison to males, the female reproductive system is not typically as useful for differentiating lower taxonomic ranks, and is not used throughout this thesis. For this reason the female reproductive system will not be detailed here, but for a very detailed explanation of this system, readers can consult McGregor and Kesling (1969a,b). Male cypridoidean ostracods typically have paired symmetrical reproductive systems (left and right) that have been reported in at least one case as being completely unconnected (McGregor and Kesling, 1969a). Sex cells de- velop within a number of testes that line the inner surface of the left and right valves posteriorly (Figure 2.11). They are often visible through the outside of the carapace with a dissecting microscope and this makes them very useful features to easily differentiate males and females. The multiple testes join to form a single conduit on each side called the vas deferens. These run through the body, then around the interior margin of each valve, starting antero- dorsally, around the ventral margin, up the posterior margin, and along the dorsal margin where it then loops multiple times. Here, the paired vas deferens widen abruptly to form large paired seminal vesicles (Figure 2.11) which occupy much of the dorsal region of the body. At the posterior end of each of the seminal vesicles, two large organs on each side called Zenker’s organs pul- sate with muscular contractions. Each pumps spermatozoa through a central tube within each Zenker’s organ, both of which then run toward a hemipenis that lies on the underside of the body. The hemipenis is usually highly variable between species (Figure 2.11), and is very useful for species identiﬁca- tion when males are present in a population sample. It is a three dimensional

2.6. REPRODUCTIVE SYSTEM 43 structure that consists of a lateral and medial shield, sometimes with additional lobes, which vary in shape and size, as do the coiling patterns of the spermiduct within. This concludes a brief introduction to the fundamental morphology of cypridoidean ostracods, and the terms used to describe them throughout this thesis. It is hoped that this will serve as a reference guide from which readers can occasionally return to if needed, for explanation of features described in the following chapters.

Figure 2.1: Terminology used in descriptive literature to indicate the position of features of the carapace and limbs in lateral (top) and dorsal (bottom) views, also showing common length measurements of the valves.

44 CHAPTER 2. THE CYPRIDOIDEA Figure 2.2: Major internal valve features for Cypridoidea, showing illustration of Ilyodromus intermedius with left and right valves dissected and body removed. Line drawings taken from Smith et al. (2011).

2.6. REPRODUCTIVE SYSTEM 45 Figure 2.3: Nomenclature for the duplicature and wall structure of a cypridoidean valve, showing cross section of a single hypothetical valve with important features exaggerated in size. Redrawn after Van Morkhoven (1962).

46 CHAPTER 2. THE CYPRIDOIDEA Figure 2.4: Schematic diagram of the typical cypridoidean appendage organisation in antero-lateral view (Ilyodromus sp.), with right valve artiﬁcially over-transparent to enable visibility of the limbs

2.6. REPRODUCTIVE SYSTEM 47 Figure 2.5: Morphology and nomenclature of the cypridoidean antennule and antenna, showing illustrations of Ilyodromus stanleyanus appendages, with segment numbering system. Line drawings taken from Shearn et al. (2014). Scales: Antennule = 200 µm, Antenna = 150 µm.

48 CHAPTER 2. THE CYPRIDOIDEA Figure 2.6: Morphology and nomenclature of the cypridoidean mandibula, showing illustrations from Chlamydotheca colombiensis, with segment numbering system and the positions of important setae. Scale = 273 µm.

2.6. REPRODUCTIVE SYSTEM 49 Figure 2.7: Morphology and nomenclature of the cypridoidean maxillula, showing illustrations from Chlamydotheca colombiensis, without detail of chaetotaxy on endites I and II. Scale: Branchial plate = 273 µm, Palp and endites = 205 µm.

50 CHAPTER 2. THE CYPRIDOIDEA Figure 2.8: Morphology and nomenclature of the cypridoidean ﬁfth limb, showing illustrations from Ilyodromus sp., segment organisation, position of setae and corresponding endopod in males. Scale = approx 200 µm.

2.6. REPRODUCTIVE SYSTEM 51 Figure 2.9: Morphology and nomenclature of the cypridoidean sixth and seventh limbs, showing illustrations from Ilyodromus stanleyanus and segment numbering system. Line drawings taken from Shearn et al. (2014). Scale = 150 µm.

52 CHAPTER 2. THE CYPRIDOIDEA Figure 2.10: Morphology and nomenclature of the cypridoidean caudal ramus and caudal ramus attachment, showing illustrations from Ilyodromus stanleyanus. Line drawings taken from Shearn et al. (2014). Scale = 150 µm.

2.6. REPRODUCTIVE SYSTEM 53 Figure 2.11: Top: Morphological nomenclature and organisation of the male cypridoidean reproductive system in right lateral view, with hole in the right valve to enable visibility of internal structures. Illustration modiﬁed from Matzke-Karasz et al. (2014b). Bottom: Vari- ability in hemipenis morphology, indicating major homologous structures. Line drawings of Bennelongia taken from Shearn et al. (2012). Drawings not to scale.

54 CHAPTER 2. THE CYPRIDOIDEA 2.7 References

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