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INLAND CRUSTACEAN FISHERIES

Miles Abadilla, W. Ray McClain, Taku Sato, Luis M. Mejía-Ortíz, and Miguel A. Penna-Díaz

Abstract Freshwater crustacean inland fisheries are typically confined to small local areas that are associated with bodies of water, like rivers and swamps. They are small-scale fisheries, enough to supply the requirements for local commercial use, and considered mainly as a subsistence resource for small fishing communities. As such, inland crustacean fisheries exemplify a subsistence opportunity for small countries with limited economic power, particularly rural communities. Crustaceans are a relevant protein source alternative to fish, and they are often also associated with local gastronomical and cultural identity. Data for the most commonly caught species are often unavailable since these fisheries exist in remote areas where government or institu- tional monitoring is limited, making it difficult to obtain reliable data about small-scale fisheries. Nonetheless, the captured species and the techniques used are widely diverse. In inland fisheries, the main target species are prawns from the genus Macrobrachium, gathered within the tropics all over the world. Natural fisheries of crayfishes in their native range exist on several continents, with Procambarus clarkii, Pacifastacus leniusculus, and Astacus astacus as the main commercial species. Some of them have1 also become a fisheries resource in areas where they were introduced. The only terrestrial crab fishery is for an anomuran, the coconut crabBirgus latro. The future of most of these fisheries depends in part on the health of the water sources threatened by con- tamination and unrestricted waterways jeopardized by the construction of dam-like structures that block the migration of some species to complete their reproductive cycles. The creation of regulatory policies is key for keeping the fishery activity self-sustainable because most of the exploitation depends on wild populations. There are some conservation efforts implemented thanks to the relevant gastronomical value of the species, as well as to economically sustain local communities in remote areas.

Miles Abadilla, W. Ray McClain, Taku Sato, Luis M. Mejía-Ortíz, and Miguel A. Penna-Díaz, Inland Crustacean Fisheries In: Fisheries and Aquaculture. Edited by: Gustavo Lovrich and Martin Thiel, Oxford University Press (2020). © Oxford University Press. DOI: 10.1093/oso/9780190865627.003.0008 182 --- Not for reuse or distribution ---

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INTRODUCTION

Crustaceans are fished not only in the marine environment but also in inland habitats such as lakes, rivers, and swamps, as well as on land, where some terrestrial crustaceans are caught for either commercial or subsistence use. The global catch data of these freshwater and semi-ter restrial inland crustaceans tend to be undocumented because most of this activity constitutes small-scale artisanal or subsistence fisheries that exploit local crustacean species (Welcomme 2011). Likewise, research on these fisheries is also largely lacking (New et al. 2000). For example, in southern central Chile, the crayfish Samastacus spinifrons is harvested in rivers and lakes through a variety of methods and then sold locally. However, no data exist on their catch since Chilean fishery regulations do not rec- ognize this species as a fishery resource (Rudolph 2013a). Similarly, in the myriad islets of the tropical Indo-Pacific region, the coconut crabBirgus latro is traditionally and locally consumed, yet little is known about the harvest sizes across the different islands and countries (Sato and Yoseda 2010). Inland crustaceans are extremely diverse, with different species, lifestyles, life histories, and reproductive biology, influencing the extent of human exploitation. Many freshwater crustaceans, such as all freshwater crabs, have direct development (i.e., there is no larval stage during development, with the embryo developing into an adult-lik e juvenile) (Strathmann 2018). Direct development limits the dispersal ability of these species relative to their counterparts that have indirect development, which includes a free-living larval stage for dispersal into other habitats. Hence, the habitat range and geographic distribution of landlocked species are also usually restricted to a specific water body or region; consequently, only local communities may encounter and harvest these crustaceans for subsistence or for small-scale commercial use. In some large-scale inland crustacean fisheries, such as the Chinese mitten crabEriocheir ( sinensis), the effect of lower natural catch sizes has been bolstered by farmed production during the final decades of the twentieth century. Currently, most, if not all, industrial-scale exploitation of inland crustaceans is dependent on aquaculture, which has largely exceeded wild capture yields to meet global demand (New et al. 2000, Sui et al. 2011). In this chapter, we explore inland terrestrial and freshwater crustacean fisheries that are economically important to the regions where these crustaceans occur. The following sections detail se- lect crustacean fisheries that are most representative of the main crustacean groups: prawns, crabs, and crayfishes. Each case study highlights the biological and ecological features that allow these organisms to be suitable for capture fisheries in their location, management practices (if any), and the future outlook for the species.

FRESHWATER PRAWNS

Prawns of the genus Macrobrachium belongs to the family Palaemonidae and are distributed mainly in tropical and subtropical regions. There are over 240 recognized species ofMacrobrachium around the world (Wowor1 et al. 2009). Approximately 190 species are diadromous, and 50 of them have direct development. The distribution limits for these species are at 35º of latitude in both hemispheres (Anger 2013), and they inhabit four continents: the Americas, Africa, Asia, and Oceania (Fig. 8.1). Their habitats are diverse since there are species living in caves, springs, streams, and rivers, and they can also be found in freshwater or coastal lagoons with variable salinities. The recent discoveries of new species in caves, groundwater systems, wetlands, or other isolated areas suggest that this group may be experiencing ongoing speciation (Komai and Fujita 2005, Dos Santos et al. 2013, Cai and Vidthayanon 2016). Prawns are common in the tropical and subtropical rivers with access to the sea. There are two main groups: the first is composed of species that can conquer new ecological niches, have fast --- Not for reuse or distribution ---

Inland Crustacean Fisheries 183

Fig. 8.1. Global records of the Macrobrachium spp. and their distribution ranges based on global records. Black dots represent coordinates and centroids of the current records for Macrobrachium spp. as of June 12, 2019. Figure modified after the map generated by Discover Life’s (2019) Global Mapper using the database of Global Biodiversity Information Facility (1,524); CNCR [Colección Nacional de Crustaceos] (730); Australian Museum provider for OZCAM (196); Rapid Assessment Program (RAP) Biodiversity Survey Database (163); NMNH Invertebrate Zoology Collections (119); Albany Museum (59); Collection Crustacea, ZMB (52); CAS Invertebrate Zoology (IZ) (50); invertebratezoology (44); Museum of Comparative Zoology, Harvard University (40); Field Museum of Natural History (Zoology) Invertebrate Collection (39); Invertebrates (GBIF-SE:SMNH) (29); Peabody Invertebrate Zoology DiGIR Service (2); Crustaceans specimens (2); Atlantic Reference Centre (OBIS Canada) (1); and iNaturalist (55). Numbers in parenthesis are records from each database.

growth, and produce many offspring, with free-living larval stages in brackish and marine waters. The larvae develop in the sea until reaching the juvenile stage. From there, juveniles move from the coastal areas and migrate upstream to rivers to grow and reproduce, making them a diadromous species (Bauer 2013). A second group involves species with direct development, which do not need brackish or seawater in their first free-living stage (Jalihal et al. 1993, Anger 2013, 2016). Diadromous prawns produce higher yields since these species are larger in size, adding to their value and attractiveness to locals. Species with direct development are generally smaller and can only be found in local markets, mainly consumed by the local indigenous people (Mejía-Ortíz et al. 2016, Tejeda-Mazariegos et al. 2018). Macrobrachium rosenbergii (Fig. 8.2) is widely cultivated because this species has an important international market, with an estimated annual production of more than 200,000 t (New 2014; see Chapters 4 and 9 in this volume). However, its introduction for aquaculture in the different rivers of the USA, Mexico, Brazil, India, Vietnam, Indonesia, and China (New 2014) has caused 1ecological problems, affecting the native and endemic fauna (Bowles et al. 2000, Pérez et al. 2003, Magalhaes et al. 2005, Silva-Oliviera et al. 2011). In order to avoid new ecological catastrophes, there are efforts to cultivate native prawn species of economic interest within their native distribution ranges. These include M. carcinus, M. tenellum, M. acanthurus, M. americanum, M. amazonicum, and M. surinamicum in the Americas; M. macrobrachion and M. vollenvhovenii in Africa; M. rosenbergii, M. nipponense, M. malcolmsonii, M. idella, and M. lamarrei in Asia (New 2005; see Chapter 9 in this volume). In natural populations, prawn yields have decreased in the last twenty years. Unfortunately, reports are from artisanal fisheries where data are of poor quality, and the lack of long-term records to identify changes and the possible causes of such decreases (for shrimp-prawn fisheries, see 184 --- Not for reuse or distribution ---

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Fig. 8.2. The giant river prawn Macrobrachium rosenbergii. (A) M. rosenbergii adult male specimen. (B) Harvested M. rosenbergii individuals from a culture. Figures are courtesy of Wikimedia Commons (under CC Creative Commons license). See a color version of this figure in the centerfold.

Chapter 4 in this volume). An alternative explanation for such sharp declines could be due to environmental issues, such as river pollution (Dutra et al. 2016) or recent dam construction for agricultural purposes or the generation of energy. In the latter case, species migrations along rivers to their reproductive natural habitats are interrupted; thus, life cycles cannot be completed, and the population is reduced (Bauer 2011).

Biology and1 Behavior/Ecology

Freshwater prawns vary in sizes, with some species growing to the dimension of a lobster, reaching over 300 mm in body length and weighing up to 1 kg, while others are similar in size to crayfish (Bowles et al. 2000).Macrobrachium rosenbergii, for example, averages maximum sizes of 320 and 250 mm for males and females, respectively (FAO 2009). Adult Macrobrachium spp. generally prefer low-saline waters. Through hyperosmoregulation, wherein their specialized cells, tissues, and organs transport ions through cell membranes to the hemolymph, freshwater prawns can tolerate brackish or freshwater conditions. This function is absent, however, during --- Not for reuse or distribution ---

Inland Crustacean Fisheries 185 the larval stages of such diadromous species, indicating an incomplete evolutionary transition (Anger 2016). For males among Macrobrachium spp., female guarding occurs before and during her premating molt (Karplus and Barki 2019). Right after the female molt, the male turns her over and transfers the spermatophore. In general, the females release mature oocytes, which are fertilized while being passed and attached to the pleopods, in a process that can last between two to three hours (Lynn and Clark 1983). During this process, the female is vulnerable to cannibalization by other males. Her mate continues to protect her from the aggression of other males and guards her to avoid sperm competition with other males (Karplus and Barki 2019). After mating, females hide in refuges to conclude their exoskeleton chitinization (Kruangkum et al. 2015). In Macrobrachium species, as in all other crustaceans, embryogenesis varies in duration between species, depending on environmental temperatures (Lewis et al. 1966, Choudhury 1971) and the habitat used by ovigerous females (Bauer and Delahoussaye 2008, Bauer 2013). For instance, female M. heterochirus, which inhabit the upper river, move downstream when it rains and the water flow increases, to hatch their eggs close to coastal areas. Thus, their larvae can reach the brackish waters quickly (Mejía-Ortíz et al. 2001). In several species, such as M. carcinus and M. acanthurus, ovigerous females living in the midreaches also migrate downstream (Mejía-Ortíz and Alvarez 2010). Cryphiops caementarius can migrate downriver as much as 100 km to go to the river mouths (Hartmann 1958). In most Macrobrachium species, larval development occurs in the marine environment; after the juvenile stage is attained, upstream migration to different areas begins, based on their specific developmental requirements (Bauer 2018). On the other hand, species that thrive exclusively in freshwater do not export larvae to brackish waters because they cannot osmoregulate to survive in seawater (Anger 2013). Some freshwater prawn species have a partially abbreviated or completely abbreviated larval development with variable times of embryogenesis (Jalihal et al. 1993, Alvarez et al. 2002, Mejía-Ortíz et al. 2010). Species with complete abbreviated larval development have low fecundity, generally between 20 and 100 large eggs, and intraspecific egg cannibalization is frequent among these species (Ordiano et al. 2005). Seasonality can affect the feeding behavior and abundance of certainMacrobrachium species as the rainy season transports more organic matter from the forest into the river (Mejía-Ortíz and Alvarez 2010). For example, in the Huitzilapan River, Mexico, the abundances of M. acanthurus and M. carcinus increase during the rainy season, whereas M. heterochirus populations thrive only during the dry season in the upper and middle reaches of the river (Mejía-Ortíz and Alvarez 2010).

Fishing Methods

A combination of tools and materials can be used for fishing, including cast nets, trident spears, small dragnets with brushwood as prawn shelter, or traps with bait (New et al. 2000). In artisanal fisheries, baited traps are commonly used for catching freshwater prawns (Table 8.1). Traps are tied to a long line, whose1 location is marked by a flagged pole (New et al. 2000). The trap design uses the principle of an inverted funnel, commonly made of bamboo or riparian vegetation roots, and some can also be made with metallic mesh (Bauer and Delahoussaye 2008, Embrose and Isangedihi 2016). In general, fishers place the traps inside ponds or rivers with a special bread, fish, or meat as bait and check them during the night. This method is used for those species that live mostly in the lower reaches of the river or within coastal zones. For other species, fishers use a cast net that closes when pulled to catch the animals. For example, fisheries in the Hooghly and Godavari rivers of India contain stranded prawns, captured in ebbing waters during the dusk and dawn (New et al. 2000). 186 --- Not for reuse or distribution ---

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Table 8.1. Macrobrachium species with an artisanal fishery in several countries.

Species Country M. australe Madagascar M. duz Zaire and Nigeria M. idai Madagascar M. idella Tanzania M. lar Mauritius M. lepidactylus Kenya, Madagascar, and Tanzania M. macrobrachium Benin, Liberia, Guinea, and Nigeria M. patsa Madagascar M. raridens Ghana and Guinea M. rude Kenya and Tanzania M. scabriculum Kenya and Tanzania M. vollenhovenii Cote d’ Ivoire, Gabon, Guinea, Liberia, Senegal, Zaire, and Nigeria M. acanthurus Brazil, Mexico, and Venezuela M. carcinus Mexico, Brazil, Honduras, Guatemala, and Venezuela M. amazonicum Brazil, Guyana, Suriname, and Venezuela M. americanum El Salvador, Guatemala, Mexico, Colombia, Costa Rica, and Peru M. heterochirus Brazil and Mexico M. jelskii Brazil M. ohione USA M. olfersii Brazil, Venezuela, and Mexico M. tenellum El Salvador and Mexico M. rosenbergii Pakistan, India, and Bangladesh M. malcolmsonii Pakistan and India M. choprai India M. rude India M. idella India M. scabriculum India M. equidens India M. vollenhovenii India

Current Management

In Mexico and Latin America, most artisanal fishing activities are related to the rainy season since the fishery1 depends on occurrence of rain (García-Guerrero et al. 2013). Unfortunately, there are no local laws to define fishing seasons since there is a general lack of biological information that would allow confidently establishing closed seasons. In most fished areas, fishery unions are nonexistent; hence, there is no established minimum catch size. However, most fishers do not harvest prawns during the reproductive season because the market rejects ovigerous females. Since fishing for prawns is seasonal and complementary to other activities, fishers supplement their prawn catch by catching other freshwater animals or by working in agriculture. All forms of prawn culture are for larger species, whereas fisheries for smaller species are generally not documented since they are mostly for self-consumption or for local and small trade. --- Not for reuse or distribution ---

Inland Crustacean Fisheries 187

While M. rosenbergi and M. malcolmsonii constitute the main freshwater prawns produced on the Indian subcontinent, there are artisanal fisheries sprinkled throughout the region. Other recorded species include the Ganges river prawn (M. choprai), the hairy river prawn (M. rude), the slender river prawn (M. idella), the Goda river prawn (M. scabriculum), and the rough river prawn (M. equidens), to name a few. Some prawn species are not exported; rather, they are consumed locally instead, despite the proximity of processing factories, like M. lamarrei in Bangladesh. Likewise, postlarvae and juveniles of M. vollenhovenii are of particular importance in the Senegal River Basin as they are used for restocking of the wild populations (Alkalay et al. 2014) in addition to its local consumption as dried or fresh food (New et al. 2000).

Cryphiops caementarius

The Andes river prawn is distributed along the rivers of southwestern Peru and northern Chile. Cryphiops caementarius is closely related to the genus Macrobrachium and, consequently, has a very similar biology (Meruane et al. 2006, Pileggi and Mantelatto. 2010). As mostMacrobrachium species, C. caementarius is sexually dimorphic. Furthermore, males present two different morphotypes: one of an average size and regular shape while the other is larger, with several strong teeth modified to damage and puncture the exoskeleton of rivals in agonistic encounters related to territorial disputes or access to females (Rojas et al. 2012). This larger morphotype is called “garrudo” by local fishers (Fig. 8.3A). Sexual maturity ranges from a minimum of 9 to 10 mm to 33 to 36 mm carapace length (CL). The spawning period is during the austral spring and summer, between September and March, with the highest proportion of ovigerous females occurring between November and January. Fecundity is size dependent, ranging from 15,000 to 36,000 eggs in females of 22 to 36 mm CL, respectively (Meruane et al. 2006, Morales et al. 2006). The egg volume and female fecundity depend directly on the female diet composition (Bazán et al. 2009). Larvae of C. caementarius can disperse in the sea, which explains the high genetic diversity and connectivity between distant river systems (Hartmann 1958, Dennenmoser et al. 2010), as reviewed by Bauer (2013). Like many of the aforementioned freshwater prawns, C. caementarius holds local commercial interest and potential for its cultivation and export. However, in-depth knowledge and records of its catch and culture are scarce. Interest for the Andean prawn was first recorded during the Spanish colonial era, which resulted in the ban of C. caementarius extraction in the present Peruvian ter- ritories because the fishing techniques used at that time drained too much water from the rivers’ courses (Viacava et al. 1978). For the Chilean territory, the government enacted a total ban of C. caementarius extraction in 1934 (Báez et al. 1983), with a most recent updated decree in 1986 that (1) bans its extraction between December and April; (2) restricts the capture of prawns below 70 mm total length or 30 mm CL between May and November; and (3) prohibits the relocation, retention, consumption or marketing of ovigerous females in all fisheries (Meruane et al. 2006, Morales and Meruane 2013). Cryphiops caementarius is neither officially recognized in the Chilean legislation as a native natural1 resource nor recognized as a national artisanal fishery in any local activity. Even the organizations directly developing its aquaculture are unofficial. Development for this fishing activity continues to grow regardless of the lack of regulation. For example, in Chile, the unemployed agricultural workers that live near the Limarí River occasionally capture juveniles from the wild and sell them to private companies that cultivate the prawns until they reach the appropriate commercial size. Later, the prawns are traded in the local market or to specialized gourmet restaurants. An adequate articulation of this commodity’s benefits to the different sectors of the local population can make this resource of economic value while also potentially improving rural economy (Morales and Meruane 2013) when there is no other employment opportunity. 188 --- Not for reuse or distribution ---

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Fig. 8.3. The Andes river prawnCryphiops caementarius. (A) Male “garrudo” morphotype with one oversize chelae to defend their territory in agonistic encounters. (B) Male adult individual hidden in their refuge. Figure 8.3A photo courtesy of Arthur Anker with permission and Figure 8.3B photo courtesy of Ivan Hinojosa with permission. See a color version of this figure in the centerfold.

Perspectives1 For the fishery of freshwater prawns of the genus Macrobrachium, the main management concerns are (1) abundances of the wild populations are decreasing, and the catch size of individuals is diminishing over time because of their overexploitation; (2) dam construction on the rivers; (3) changes in the water level of rivers due to deforestation; and (4) the increase of wastewater from cities. The first problem of overfishing is attributed to a lack of restrictions, a defined capture quota, or seasonal closure. There are, however, many fishers that observe a minimum size for retaining -an imals and release caught ovigerous females. Despite this, depending on their social and economic situation, fishers are fishing without any regulatory framework. At a local level, artisanal fisheries --- Not for reuse or distribution ---

Inland Crustacean Fisheries 189 should be regulated by considering the different actors and their social and economic conditions as well as the life histories of the exploited species. All this should aim to balance sustainable fishery practices while still benefitting fishers. The second problem is driven by energy need since many dam constructions aim to provide electricity to growing urban areas. For certain prawn species, the environmental conditions for reproduction or growing are very specific and can change based on the river’s conditions. Therefore, the building of structures that interrupt the natural river flow should consider ways to allow prawn migrations to complete their natural life cycle. The third problem is a consequence of deforestation in upper reaches (where one principal function of the forest is the retention of rainwater), causing flooding events and thus a constant decrease in the water levels of the rivers, along with the creation of newly formed river structures that become barriers for migration. Finally, the constant flow of wastewater from the cities in the river basins should be resolved. The inefficient regulation of wastewater creates negative environmental conditions that impact the habitats and ultimately the fisheries for these species. Freshwater prawns are exploited through local fisheries in different areas around the world since they are valued based on their varied size, flavor, and price. However, these fisheries have several concerns, and they will benefit from proper regulation and constant care to achieve a sustainable activity. The main points discussed offer a good opportunity to produce scientific information that can be used in the management of these natural resources, such as differences in population dy- namics, decreases in the size at catch, and changes in the reproductive areas.

TERRESTRIAL HERMIT CRABS

Terrestrial hermit crabs belong to the family Coenobitidae (Anomura), which includes two genera, the genus Coenobita with 16 species of the land hermit crabs and the genus Birgus, with only one species, the coconut crab Birgus latro (McLaughlin et al. 2010). The following sections focus on the latter.

Biology of the Coconut Crab

The coconut crab, also known as robber crab or palm thief, is the largest living terrestrial arthropod, can weigh up to 4 kg (Brown and Fielder 1991), and lives approximately 50 years (Sato et al. 2013, Oka et al. 2015). Although coconut crab juveniles retain the shell-carrying habit of its hermit crab ancestors (Reese 1968), adults have no gastropod shell, and their abdomen is protected on the dorsal surface by a series of hardened tergal plates. Although this species is fully terrestrial, individuals must go regularly to the sea to ingest seawater in order to maintain their osmotic balance (Gross 1955). Furthermore, females must return to the sea to hatch their eggs1 (Schiller et al. 1991). Hatched larvae develop through several planktonic zoeal stages in the ocean before they metamorphose into a benthic megalopa (Reese and Kinzie 1968). After settlement, megalopae grab and carry gastropod shells, and then migrate to land (e.g., Hamasaki et al. 2015b). Through such dispersal and recruitment processes, their geographical distribution and population connectivity are determined. Coconut crabs inhabit oceanic islets and atolls as well as islands in the tropical Indo-Pacific coastal region. Their distributional range includes islands off the eastern coast of Africa near Zanzibar in Tanzania and eastward to the Gambier Islands, French Polynesia, in the east Pacific (Drew et al. 2010). Lavery et al. (1996) inferred that coconut crabs colonized the Pacific region from the Indian Ocean. In the northwestern Pacific region, coconut crabs experienced population expansion and rapidly colonized there during the glacial 190 --- Not for reuse or distribution ---

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period (Hamasaki et al. 2015a). Subtropical populations also occur in the Ryukyu Islands, Japan, and Taiwan (Drew et al. 2010). Coconut crabs can be sexed according to the presence of pleopods on the left ventral surface: only females have pleopods to support their eggs externally (Fletcher 1993). They exhibit sex-specific growth patterns, with males growing faster than females, and a pronounced sexual size dimorphism in which the mean and asymptotic body size of males is larger than those of females (Sato et al. 2013, Oka et al. 2015). Sexually mature individuals show a clear sexual dimorphism, with males having considerably larger chelipeds than females (Anagnostou and Schubart 2014). These sexual differences are attributed to sexual selection processes and energetic costs of reproduction (Sato et al. 2013, Anagnostou and Schubart 2014). Body size is commonly expressed in thoracic length (hereafter, ThL). Females attain functional maturity at 24.5 mm ThL (Sato and Yoseda 2008) and at approximately nine years of age (Sato et al. 2013). Males >25 mm ThL are considered physiologically mature (Sato et al. 2008) and at ~6 years old (Sato et al. 2013).

Reproductive Ecology of the Coconut Crab

Recently, several aspects of the reproduction of coconut crabs gradually came to be known. Reproductive season varies throughout their distributional range (Drew et al. 2010). During the reproductive season, both males and females migrate from inland areas to the coast for the pur- pose of acquiring mates and larval release into the ocean, respectively (Sato and Yoseda 2013). Males hold many spermatophores containing a great quantity of sperm in the vasa deferentia (Sato et al. 2008). Mating is thought to occur near the seashore (Sato and Yoseda 2013) around the new moon phase (Sato and Yoseda 2009b), but the exact timing has not been determined (Drew et al. 2010). Mating is not accompanied by pre- or postcopulatory guarding behavior (Schiller et al. 1991). Female pleonal expansion is strongly correlated with female reproductive activity, and the index of pleonal expansion can be used as a morphological criterion for selecting females that are able to mate (Sato and Yoseda 2009b). During mating, males deposit spermatophore masses over and near the gonopores on the female ventral surface. Mated females extrude eggs that are fertilized externally, within the brood chamber formed by a pleon, in burrows, near the seashore (Sato and Yoseda 2009a). The time lag between mating and egg extrusion is estimated to be less than one week (Sato and Yoseda 2009b). Females extrude only one clutch in a reproductive season (Sato and Yoseda 2008). There is a positive correlation between female body size and the number of eggs extruded (approximately 50,000–250,000 eggs) (Sato and Yoseda 2008). Embryogenesis is estimated to be 27–29 days (Schiller et al. 1991), during which females remain near the seashore (Sato and Yoseda 2013) due to seawater requirements for egg incubation (Schiller et al. 1991). Although larval release into the ocean is thought to be associated with lunar and tidal rhythms (Schiller et al. 1991, Sato and Yoseda 2009b, Drew et al. 2010), the exact timing has been contested. Current Status1 and Management Measures Due to its large body size and palatability of its meat, coconut crabs have been traditionally eaten by locals and are an economically important food and revenue resource for locals. Coconut crabs are easily harvested by hand and require no specialized equipment and investment because they live a purely terrestrial existence scavenging in the forest and on the seashore and feature a slow walking speed (0.15 km h-1) (Krieger et al. 2012). Furthermore, the aggregation habits of coconut crabs facilitate their harvesting. Coconut crabs can readily locate and be attracted to smelly foods (e.g., carrion of any sort, fruits, and pith of fallen trees) (Stensmyr et al. 2005), forming large feeding congregations around food sources (Drew and Hansson 2014, Krieger et al. 2016). Locals and hunters often bait the trap with, for example, coconut and flesh to catch them efficiently. --- Not for reuse or distribution ---

Inland Crustacean Fisheries 191

Currently, populations in most habitats have been severely depleted (Brown and Fielder 1991) due primarily to overharvesting (Fletcher 1993) and habitat destruction (Eldredge 1996). In 1981, the IUCN Red List listed the coconut crab as a vulnerable species. In 1996, the listing was downgraded to the data-deficient category, not because the species had recovered but because of the lack of reliable biological data (Eldredge 1996). Recently, crabs have been harvested not only for subsistence reasons but also for commercial purposes. For example, coconut crab dishes, steamed or boiled muscle and hepatopancreas, are served for tourists in some restaurants and bars as a food specialty of Okinawa at Ishigaki Island, one island of the Sakishima archipelago (Sato and Yoseda 2010). To meet the rapidly increasing demand for commercial purposes, crabs are harvested and shipped from several islands within the archipelago. In Vanuatu, for example, harvested crabs are sent by air from adjacent small islands to cover the touristic demand (Shokita 2011). Various management measures for coconut crabs have been used or suggested in several areas (e.g., Guam, Vanuatu, Commonwealth of Northern Mariana Islands, Federated States of Micronesia, Tuvalu, Solomon Islands, and Christmas Island) (Fletcher 1993, Kessler 2006, Drew et al. 2010, Buden 2012) as follows: (1) minimum size limit, (2) prohibition on harvesting ovigerous females, (3) closed season coinciding with reproductive season or lunar cycle, (4) catch limit (bag or season limits), (5) sanctuaries where harvesting is restricted, and (6) prohibition on commercial export and trade. However, most of these limits were formulated without detailed knowledge of the biology and ecology of this crab in most regions (Brown and Fielder 1991, Drew et al. 2010, Buden 2012). The lack of information on the reproductive ecology of this species hinders our understanding of how current harvesting influences the resource and has resulted in undesired consequences (Sato 2012).

Undesired Consequences Due to Current Management

The sexual size dimorphism can cause sex-biased harvesting. In species in which males grow to a much larger size than females, minimum size limits will result in selective harvest of large males. In some regions, minimum size limits are implemented (e.g., Vanuatu or in the Sakishima archipelago), and large coconut crab males are selectively harvested and commercialized (Fig. 8.4A; Sato and Yoseda 2010, Shokita 2011). In these harvested populations, mean male size decreases, and the sex ratio skews toward females (Fig. 8.4B; Sato and Yoseda 2010). In such harvested populations, small males replace large males in reproduction, and the few remaining large males participate in more matings than in pristine populations. Coconut crab males have size-dependent reproductive potentials. Larger males can achieve more matings than smaller ones (Sato 2011), as larger males have larger sperm reserves than smaller ones (Sato et al. 2008). Small males provide females with fewer spermatozoids than larger males do, which results in lower fertilization rates due to sperm limitation (Sato et al. 2010). The number of ejaculated sperm decreases with increasing male mating frequency, regardless of male size, which1 is attributed to slow recovery rate of sperm reserves (Sato et al. 2010). However, larger males always transfer more sperm to successive mates than smaller ones (Sato et al. 2010). Therefore, the sperm availability decreases sharply in large male-selective harvested populations, which consist of small males and a female-biased sex ratio (Sato 2011). In harvested populations on Hatoma Island, more than half of the females retain very low numbers of sperm, which will limit their fertilization rates (Sato et al. 2010); the remaining males mate successively and deplete their sperm reserves to the point of sperm exhaustion when they are no longer able to mate (Sato 2011). In large male-selective harvested populations, reproductive output can be affected by availability of both sperm and males (Sato 2012). Coconut crab females exhibit mate preference for 192 --- Not for reuse or distribution ---

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(A) 250

200 Male Female 150

100

No. of individuals 50

(B) 250 Male 200 Female

150

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No. of individuals 50

0 0–10 10–15 15–20 20–25 25–30 30–35 35–40 40–45 45–50 50–55 55–60 60–65 65–70 70–75 75–80 Thoracic length (mm)

Fig. 8.4. Size frequency distributions by sex of (A) coconut crabs marketed for consumption in Ishigaki Island in 2008 (male, n = 1,055; female, n = 50); and (B) individuals in a harvested population at Hatoma Island in 2007 (male, n = 453; female, n = 557). Redrawn after Sato and Yoseda 2010.

larger males, and they only mate with males larger than or similar in size to themselves (Sato and Yoseda 2010). A decrease in the density of large males due to harvesting reduces the frequency of female encounters with favorite mates and thus causes difficulties for females to find mates. Some females might fail to meet preferred mates within their receptive period, possibly com- promising their reproductive potential. The encounter rate between females and favored mates would be low, especially for large females, due to their mate preference. Large females will make a larger contribution to the reproductive rate and recruitment of harvested resources because (1) fecundity increases exponentially with female body size (Sato and Yoseda 2008), and (2) large females produce better larvae with respect to susceptibilities to predation and starvation (Sato and Suzuki1 2010). Coconut crab females are susceptible to the decline in sperm and mate availabilities because they have no seminal receptacles. Current large male-selective harvesting can decrease reproductive outputs of coconut crab populations through decreasing both sperm and mate availabilities, which would eventually influence abundance and stability of the resource (Fig. 8.5). Knowledge about the details of reproductive ecology of the coconut crab is surely helpful for understanding how current harvesting influences their reproductive output and its susceptibility to variation of availabilities of sperm and mate. The most logical management strategy for the protection of larger males would be multiprong management strategies that consist of, for example, a sanctuary and size limits outside of this area (Sato 2012). --- Not for reuse or distribution --- Maternal inﬂuences Female mate choice Loss of mating opportunity Decrease in mate availability for females increase in male mating frequency decrease in frequency of female encounters with males Decrease of potent males Reproduction 2) Skew of sex ratio toward females Decreases in larval qualities 2012. Sato after Redrawn resources. crab coconut of abundance outputand reproductive harvesting influences selective number of retained sperm possible number of mating number of ejaculated sperm fertilization rate Decreases in reproductive output Decrease in sperm availability for females Sperm limitation Slow recovery of sperm Male size-dependent reproductive potentials

1) Decrease in mean male size 1 Change of population demographic structure Large male-selective harvesting Decreases in reproductive output and abundance of the resources Fig. 8.5. male- large of how Schema 194 --- Not for reuse or distribution ---

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Toward the Future

On Christmas Island, coconut crabs have been protected within a no-take sanctuary on over 60% of the island since 1978, resulting in a population currently thought to be one of the largest in the world (Drew et al. 2010). However, even a sanctuary has limits wherein it cannot completely protect highly mobile species such as the coconut crab (Krieger et al. 2012). Recently, male-only harvesting with size limits, sanctuaries, and seasonal closure have been implemented for coconut crabs in Ishigaki Island. These management measures are determined based on scientific data on (1) the previously mentioned reproductive ecology, (2) sex differences and seasonal variations in meat yield and bi- ochemical composition (Sato et al. 2015a,b), and (3) effects of long-term frozen storage on the chemical compositions (Sato et al. 2016). However, currently there are no data to judge which size sanctuaries is appropriate for this species. To determine the appropriate size that will provide them with effective and long-ter m protection, further study about their long-ter m movement patterns is needed (though short-ter m patterns have been observed; see Krieger et al. 2012). In addition to regulations on harvesting, we must develop education programs explaining, for example, the current status of coconut crabs; vulnerability to excessive harvesting due to their life history characteristics (e.g., slow growth rate and late reproduction); the importance of habitat conservation for their food sources and reproduction; and the impacts of exotic species on coconut crab populations.

NATURAL CRAYFISH FISHERIES

Crayfish have been exploited by humankind largely wherever they were found. They have been the subject of subsistence, recreational, or commercial fisheries in several regions of the world, dating back centuries in some cases. Three families of crayfish, encompassing more than 500 species, occupy a wide range of habitat types spanning the globe. North America hosts 77% of the world’s crayfish species, including 99% of the members of the family Cambaridae as well as a single genus of the family Astacidae (Taylor, 2002). Australasia is home to a diverse 20% of the species, all in the family Parastacidae, while approximately 1.5% of the world’s species are found in South America (Parastacidae) and 1.5% are found in Eurasia (Astacidae). Most of the demand for crayfish is for human food, although some are sold for fish bait, pets, or as scientific specimens. Crayfish for food are marketed as live animals, whole prepared product, or processed abdominal meat. In parts of Europe and North America, crayfish have also been closely aligned with certain cultural traditions (Huner 1994). Aside from natural fisheries, crayfish have also been the focus of aquaculture endeavors beginning at least as far back as the sixteenth century in Europe (Ackefors and Lindqvist 1994). The history of crayfish aquaculture is intrinsically linked to the natural fishery and was originally employed to increase supply for markets or as a means of enhancing or restoring the natural fishery of a region. Crayfish aquaculture is covered in Chapter 11 in this volume.

Taxa of Commercial Interests1 While crayfishes represent dominant, keystone benthic invertebrates in many aquatic ecosystems, fewer than two dozen crayfish species currently sustain commercial fisheries. These fisheries are located principally in the USA, Spain, Scandinavia, Turkey, Australia, and mainland China, though small-scale endeavors can be found in other parts of Europe, Asia, North and South America, and Africa. The red swamp crayfishProcambarus ( clarkii; family Cambaridae), a native of southern-central USA and northeastern Mexico, accounts for approximately 70–80% of the total crayfish harvested worldwide. This is due to a thriving capture fishery in Louisiana, a state in the southern USA, and the fact that P. clarkii has been introduced widely outside of its native range, where populations in some --- Not for reuse or distribution ---

Inland Crustacean Fisheries 195 regions have ballooned to levels suitable for exploitation. Introductions in Spain, Africa, and China have led to commercial fisheries, and several locations within the United States, Europe, and Japan have populations of P. clarkii that support viable recreational fisheries (Loureiro et al. 2015). Natural fisheries of other crayfishes in their native range exist on several continents. The most valued species in Europe is the noble crayfish Astacus( astacus). It is found primarily in central, western, and northern Europe, most notably Scandinavia. However, the total annual yield of noble crayfish is only about 5% of the amount achieved prior to the introduction of the crayfish plague (Aphanomyces astaci) in 1860, which devastated astacid populations all over Europe (Skurdal and Taugbol 2002). The narrow-clawed crayfish Astacus( leptodactylus) is another important crayfish, harvested primarily from Eastern Europe, Turkey, and other countries in the Caspian region of Asia. Three other European species are recognized, Austropotamobius torrentium, A. pallipes, and Astacus pachypus, but these have less commercial significance in terms of total value. Australia is home to a diverse nine genera of crayfish in the family Parastacidae, but only three have been the subject of recreational fishing: Euastacus, Astacopsis, and Cherax. The only species for which a commercial fishery developed is the yabby Cherax( destructor). Overharvesting and ad- verse environmental conditions ultimately limited the commercial fishery, and conservation threats have resulted in increased regulations governing both recreational and commercial fisheries (Mills et al. 1994). The red claw crayfishC. quadricarinatus ( ) is a native tropical Australian species that has been widely introduced outside of Australia for aquaculture, and as a result, harvestable quantities of red claw from wild populations have been reported from Ecuador (Policar and Kazak 2015) and Mexico (Vega-Villasante et al. 2015). Native and translocated populations of the North American genus Orconectes (family Cambaridae) are also harvested in central and northern USA, although mainly for fish bait. Procambarus zonangulus occupy much of the same habitat and are captured alongside P. clarkii in the southern USA, although at much lower numbers. A commercial fishery for another native American species, the signal crayfish (Pacifastacus leniusculus) occurs along the West Coast of the USA, although the scale of this fishery is diminutive compared to the Louisiana crayfish fishery. The signal crayfish, a plague-resistant species, was also introduced to Europe beginning in the 1960s, with little foresight of the ecological implications, to replace the plague-decimated native astacid populations. Populations of the signal crayfish thrived and outcompeted the indigenous species to the point of sustaining profitable fisheries that have steadily increased since the introductions (Lewis 2002). In South America, semiterrestrial species of the genus Parastacus are widely distributed in Brazil, Paraguay, Uruguay, Argentina, and Chile. Small-scale fisheries of local species occur throughout the region, and it is estimated that up to 50 million individuals of P. pugnax are extracted annually in central Chile (Rudolph 2013b). Catch is mostly commercialized as live individuals on nearby markets (Rudolph 2013b). Parastacus pugnax is also of scientific interest because parents provide extended parental care for their offspring in their burrows (Palaoro et al. 2016), making it a potential candidate for future aquaculture projects (Ibarra and Arana 2012).

Scope and Magnitude1 of the Fisheries The magnitude and scale of the capture fisheries for crayfish are as diverse as the taxa and geographic regions described previously. The delineation between recreational and commercial fisheries is often blurred. Descriptors of the participants in any fishery scenario can range from hobbyists, intent on acquiring small quantities of crayfish for personal consumption, to those fully engaged as their primary means of employment. However, even for the most commercialized crayfishing segments, participation is predicated on individual efforts. Harvesting is not achieved by large corporations operating sophisticated equipment, but rather by collections of individuals employing simple gear. 196 --- Not for reuse or distribution ---

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Commercial fisheries for crayfish generally operate on a seasonal basis, and production can vary widely from year to year; longer trends in variations are not uncommon. Due to the nature and variability of the different fisheries and the various tendencies of regulatory agencies involved, the size and scope of the crayfish fishery globally are poorly documented. Likewise, corresponding production data for the capture fishery is woefully inadequate, largely because the measures for data collection are either nonexistent or the means to differentiate yields between the capture fishery and aquaculture are not in place. Policar and Kazak (2015) provided an excellent summary, to the extent possible, of estimated fisheries data from the various regions and countries. Because the red swamp crayfish P. clarkii( ) dominates the capture fishery of crayfish worldwide, this section is largely devoted to this species. In its native range, the largest fishery occurs in the southern (USA) state of Louisiana, where annual production of procambarid crayfish ranges from approximately 4,000 to 20,000 t (see Figure 11.1 in Chapter 11 in this volume) and contributes in ex- cess of several million US dollars to the state’s economy annually. Another 1,500 t of feral P. clarkii are captured and sold from Spain and bordering areas of Portugal, and while yields in China from aquaculture versus the capture fishery are not differentiated, it has been reported that over 479,000 t of red swamp crayfish are produced from mainland China annually (Policar and Kazak 2015). As a comparison, these authors indicated annual yields from all crayfish (both native and introduced species and from both farmed and wild-caught sources) in other Asian and European countries total approximately 277,000 and 281,000 t, respectively.

Biology of Commercial Crayfish

All commercial fisheries of crayfish involve temperate species, and most have the potential to grow relatively large (50–80 g or larger), though some dwarf species (Cambarellus spp.) are consumed in Mexico. As with all Crustacea, crayfish growth involves periodic episodes of molting with an intermolt period between each shedding incident. Molting frequency, relative molt increment, and number of molts to maturity differ among species and are largely governed by environmental conditions. Different species have different abilities to cope with various water quality conditions, diseases, and other stressors (Holdich 2002). While most crayfish have the ability to burrow somewhat, the commercial species vary considerably in their propensity for burrowing and in the types of burrows constructed. Spawning requirements also differ considerably among the taxa, and species differ with regard to habitat preferences and tolerances and to a lesser degree, food preferences. Crayfishes were originally classified primarily as herbivores, detritivores, or omnivores but are now generally recognized as obligate carnivores and facultative detritivores-herb ivores. While they will ingest myriad vascular plants, seeds, algae, vegetal detritus, and other sorted organic debris and may be able to sustain a modicum of growth from these resources, it is generally accepted that maximum growth potential must be met by balancing the diet with animal sources, typically invertebrates found in the aquatic environment (McClain et al. 1992, Momot 1995). Reproduction has been well documented for the major commercial species and differs largely by family. Age1 and size at maturity are quite variable, and mating behavior and timing differ somewhat among the species. For all commercial species, the sexes are separate (dioecious), and sexual dimorphism is exhibited at maturity. The interval between copulation and ovulation varies by species, ranging from a few days to several months in the burrowing species. Incubation period is also highly variable and largely temperature dependent. Fecundity is somewhat species specific and is also dependent on size of the female. Crayfish have fewer and larger eggs than saltwater Crustacea, whose offspring must undergo metamorphosis. Crayfish young progress through two instar stages while attached to the female and then are freed at the third instar, where they clearly resemble the adult. Two monographs (Holdich 2002, Kazak et al. 2015) provide detailed and thorough overviews of the biology of the crayfish, including comparisons among the various commercial species discussed --- Not for reuse or distribution ---

Inland Crustacean Fisheries 197 in this chapter. Specific aspects of the biology unique to the major commercial species are noted in the following material.

Procambarus clarkii

The commercial catches of crayfish in Louisiana, from both the capture fishery and aquaculture, consist of both P. clarkii and P. zonangulus, a closely related species, though P. clarkii clearly dominates. The ranges of these two species overlap within Louisiana, and they have similar ecological requirements; thus, they are often found in the same habitat. While populations of either species can be found in various locations outside of Louisiana, the largest and most important crayfish industry in North America is located largely in south central Louisiana in what is known as the Atchafalaya River Basin. This expansive (200 km long by 32 km wide) lowland area is managed as an overflow basin of the Mississippi River to the Gulf of Mexico and receives 30% of the Mississippi’s flow. The red swamp crayfishP. clarkii ( ) flourishes in the alluvial, riparian habitat of this flood- plain, characterized by seasonal flooding and drying, with the dry period usually occurring from summer through autumn. Sustained periods of river overflow permit crayfish to feed, grow, and mature. Temporary periods of dewatering promote aeration of the bottom sediments, reduce the abundance of aquatic predators, and allow for establishment of vegetation, which serves as cover for crayfish and food resources when flooded. Crayfish survive the dry intervals by digging or retreating into simple shallow burrows where they can avoid predators and acquire the moisture necessary for survival. Crayfish have also adapted to reproduce within the protection of the burrow. While P. clarkii is not a seasonal spawner in Louisiana and can spawn any time of year at that latitude, reproduction is somewhat synchronized as a result of the extended dry period, with several waves of recruits occurring from late summer to early winter (Fig. 8.6A). Red swamp crayfish is the preferred species in the southern USA marketplace because it is favored for its deep red color when cooked and its flavorful hepatopancreas (digestive gland), which is often eaten. Nonetheless, little attempt is made to separate the procambarid species for market, and no preference is shown regarding species for the packaged meat market. Red swamp crayfish are short-lived (two years or less), have high juvenile survival, and can alternate between sexually active (Form I) and inactive (Form II) stages. Growth rate is governed largely by water temperature, but acceptable harvest size (20–30 g minimum) is typically reached within a two- to four-month window. Total harvest period in Louisiana usually lasts only from about two to five months of the year, beginning as early as February or March. Incubation period is approximately three weeks in the southern USA, and fecundity rate for P. clarkii is size dependent, with brood sizes ranging from about 200–300 to well over 750 for larger specimens. Harvest of P. clarkii in Spain/Portugal and in China typically occurs during the summer months, where water temperatures are conducive for rapid growth. Yields from the capture fishery in both of these regions are influenced considerably by annual precipitation. As in Louisiana, yields are higher during rainy years and lower during low rainfall years. Serious disease problems in procambarid populations1 are rarely encountered. White spot syndrome virus has been recently found (Baumgartner et al. 2009) and may pose the greatest risk; however, no large-scale epidemic has yet been reported among wild populations of crayfish (for crustacean diseases, see Chapter 15 in this volume).

Pacifastacus leniusculus

Harvest of the signal crayfish Pacifastacus( leniusculus) from its native range in the northwestern region of North America from the 1800s until the mid-1950s occurred in lotic waters of the region (Fig. 8.6B). As more lakes and reservoirs were developed and populations were translocated in the 198 --- Not for reuse or distribution ---

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(A)

(B)

Fig. 8.6. Crayfishes. (A) Louisiana red swamp crayfishProcambarus clarkii and (B) signal crayfishPacifastacus leniusculus. Figures are courtesy of Wikimedia Commons (under CC Creative Commons license). See a color version of this figure in the centerfold.

later 1900s, commercial harvests from lentic environments emerged. The largest commercial catch of P. leniusculus1 (approximately 100,000 kg annually) occurred during the 1970s and 1980s from an introduced population in the Sacramento River Delta. Yields from the Sacramento River have waned in recent decades, but harvests of the signal crayfish in European countries, mainly Finland and Sweden, are steadily increasing (Lewis 2002). Spawning in the signal crayfish is regulated largely by photoperiod, typically occurring in October over much of its range. Incubation and hatching is governed by water temperature and can range from 166 to 280 days. Age to maturity ranges from one to three years, depending on water temperature, food resources, and crayfish density (Lewis 2002). Population density in a given habitat is generally influenced by substratum as well as availability of food (Shimizu and Goldman 1983). Signal crayfish seem to prefer rocky substrata, and though once thought not to --- Not for reuse or distribution ---

Inland Crustacean Fisheries 199 burrow, it has since been documented that they do construct simple straight or angular burrows (Guan 1994). Introductions of P. leniusculus into Europe as a replacement for the plague-decimated astacid populations further exacerbated the displacement of native species not only from a disease vector perspective, but also from excessive competition by the invasive species for shelter, food, and reproductive success (Soderback 1994). Due to its hardiness in a wide range of environmental conditions, signal crayfish are increasing all over Europe despite the general lack of management regulations, such as size restrictions or fishing seasons, to ensure its sustainability (Lewis 2002).

European Astacidae

Both A. astacus and A. leptodactylus are distributed over most of Europe, with the exception of the Iberian Peninsula, and are highly prized for their large size and preference in the marketplace. The noble crayfish A. astacus( ) can reach sizes of up to 170 mm total length (270 g) but seldom becomes larger than 150 mm. The narrow-clawed crayfish (A. leptodactylus) typically reach about 150 mm, but specimens over 300 mm also have been reported. These are cold-wat er species with similar life histories. Mating occurs in autumn, and egg development, which is temperature dependent, may take up to eight months in northern latitudes. Egg number, as with other crayfish species, is posi- tively correlated to body size of the female; however, fecundity as a function of body size is far less in the astacids than in Procambarus. Age at maturity is between three and five years (Skurdal and Taugbol 2002). Both of these species exploit a variety of habitats, such as lakes, rivers, and brooks, but the narrow-clawed crayfish can also be found in some brackish-wat er environments and more eu- trophic waters than the noble crayfish. Both prefer substrata that offer hiding places, and although they are not typically known as burrowers, they can both make small, simple burrows (Skurdal and Taugbol 2002). The crayfish plagueAphanomyces ( astaci) has represented the greatest disease risk to wild populations of Astacidae.

Harvest Gear

Harvesting of crayfish in Louisiana begins when sufficient young-of-the-year reach harvest size and when backwaters of the river basin are deep enough to allow access by commercial fishers in small boats. Typically, this occurs by spring in most years. Individual crayfish trappers in pointed-bow, flat-bottom aluminum skiffs traverse the myriad waterways to make their way deep into the inundated wooded or semi-open wetlands. Some may carry smaller paddleboats (called pirogues) to reach still deeper into the swamps. Simple, baited, wire-mesh traps spaced every 15 to 45 m are employed in lines, often running in general parallel fashion or radiating from a central location. A trapper may have multiple set locations, often some distance from one another. Traps are constructed1 from plastic-coated 1.9-cm square or hexagonal wire mesh and are made by forming a rectangular section of wire into a slightly flattened cylindrical shape and closing the ends by bringing two sides together to form a seam. A single or double funnel is formed at one end by inverting the opening at one or both corners inward. The sides of the opposite end of the pillow-shaped trap are fully or partially clamped together to form a closeable opening. “Pillow trap” dimensions vary by using wire sections of different lengths and width; however, two trap sizes are most common, 76 × 91 cm and 76 × 137 cm. Harvesters generally flatten the wire mesh traps so that they can be stacked, stored, and transported more easily, and they are simply re-formed on use. These traps are typically fished fully submerged, unlike the emergent trap styles used in shallow- water aquaculture ponds (see Figures 11.7 and 11.8A,B respectively, in Chapter 11 in this volume). 200 --- Not for reuse or distribution ---

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However, in shallow, borderline hypoxic waters, the traps can be fished in a somewhat vertical po- sition with the closed end propped up at the surface to allow trapped crayfish access to the surface. Baited lift nets, constructed of a single square piece of 1.3-cm mesh net affixed to a wire A frame at each corner, are sometimes used in shallow water by recreational fishers in the USA. The frame sits about 60 cm high and has an attachment point at its apex where it can be quickly lifted by an extension pole after a requisite soak. This type of gear is also one of several types used to harvest P. clarkii in China. Other methods used to harvest feral crayfish in China and Spain/Portugal may include homemade wire mesh traps, hoop nets, and various styles of commercially made crayfish traps, such as those produced in Europe and Australia. Although several trap types and designs are used in Europe, the main types are basically of either an upright design with the entrance on top or the prone trap with a side entrance. These traps are made from either plastic or netting and come in various configurations, often with two funneled openings. Similar traps are used for yabby in Australia. Kazak et al. (2015) provided an excellent overview and visuals of various recreational and commercial crayfish trap designs used in the various fisheries worldwide. Regardless of type or design, all traps require bait for efficient operation, with the exception of certain hoop nets (often winged) that operate via water flow. Placed in irrigation canals with the opening downstream, hoop nets are sometimes used in Spain to collect crayfish as they move upstream in the current. Bait for crayfish traps or lift nets usually consist of rough-cut fish, but other baits, such as beef spleen, poultry necks, or punctured cans of dog or cat food, are sometimes used. The commercial fishery in Louisiana utilizes formulated manufactured dry baits effectively, but only when water temperatures are above about 21°C.

Regulations

Licensing and regulations are the norm for all commercial crayfish fisheries, but requirements differ by governing entity, taxa, location, and whether a species is native or introduced. In the natural waters of Louisiana, there are no restrictions on the number of traps, length of traps, trap tags, trap removals, natural or artificial baits, minimum or maximum size of crayfish, or sex for crayfish taken under a commercial license. There are no total catch quotas, daily take, or possession limits and no restrictions on fishing seasons or day or night fishing. The only restriction currently in place is that of minimum mesh opening size for traps, which is 1.9 cm. In contrast, for Pacifastacus harvested commercially in the western USA, there is a minimum size restriction of 92 mm in the total length (from the acumen to the tip of the telson), measured in a straight line ventral side up (Thomson-Reuters Westlaw 2019). There is also either a fishing season (typically April 1 to October 31) in place or requirement to immediately return ovigerous females to the water. In Europe, fishing seasons and size limits typically apply only for native species, but minimum size limits for P. leniusculus occur in Denmark, Sweden, and France, and are 9, 10, and 12 cm, respectively (Lewis 2002). Harvest1 regulations for native commercial species in Europe are diverse and usually include restrictions on species, sex, season, size, often daily take limits, and sometimes harvesting effort or gear type. Skurdal and Taugbol (2002) provided an excellent accounting of national and fed- eral regulations for astacids, but in general the minimum size limits range from 70 to 150 mm total length. The main objective for management of crayfish in Europe is to increase stocks of native species, which principally emphasizes the control of the crayfish plague, as well as improvement and restoration of waters containing crayfish. To this end, strict regulations exist regarding the handling and translocation of crayfish. It is almost universally recognized that introductions of nonnative crayfish are not desirable and can result in unintended consequences, and contrary to just a few years ago, regulations governing introductions are now widely implemented worldwide. --- Not for reuse or distribution ---

Inland Crustacean Fisheries 201

Future Perspectives

While P. clarkii, and to a great extent P. leniusculus, are thriving within their native ranges as well as dominating and expanding well outside of their native ranges, often to the extent of threatening sensitive ecosystems, other commercial species have not fared as well. A decline in the distribution and abundance of astacids, especially the noble crayfish, due principally to the plague, often spread by an expansion of nonnative crayfish, has been devastating to that fishery. Fortunately, interest in conserving and managing native stocks and containing or slowing the spread of nonnative crayfish has increased among astacologists and governments, but the ability to effect a reversal in trend among some stocks is dubious.

SUMMARY AND CONCLUSIONS

There is a wide variety of inland crustacean fisheries, targeting different species and environments. Many of these crustaceans have been subject to aquaculture, which multiplied their yields. The fisheries remained modest, with a good degree of local interest. Here, we characterized different fisheries without historic records, poor in data and regulations such as those for freshwater prawns; the traditional fishery for the coconut crab developed in a vast territory of tropical islands, affected by a population sperm depletion from the male-only harvesting; and the better known populations with clear harvesting directives such as crayfishes. The development of established methodologies for freshwater prawns is underway, as proven by the progress made by civil society, researchers, and private organizations alike. Coconut crabs and crayfish, on the other hand, prove to be more established in the local culture, with considerable yields and marketing with significant contribution to local and regional economy. Nevertheless, in some cases regulations and fishing records need improving, which, along with further research, will result in more adequate and improved management rules. Thus, these underdeveloped fisheries can grow controlled through generating a history of records and data, continuing their sustainable consumption, and improving the livelihood of the community.

ACKNOWLEDGMENTS

We thank Mika Mei Jia Tan for her help in the planning of early versions of this chapter and putting authors in contact to make this contribution possible.

REFERENCES

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Fig. 8.3. The Andes river prawnCryphiops caementarius. (A) Male “garrudo” morphotype with one oversize chelae to defend their territory in agonistic encounters. (B) Male adult individual hidden in their refuge. Figure 8.3A photo courtesy of Arthur Anker with permission and Figure 8.3B photo courtesy of Ivan Hinojosa with permission. (A)

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