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Flowering and Seed Production in the Subtropical Seagrass, Halodule Wrightii (Shoal Grass)

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Flowering and Seed Production in the Subtropical Seagrass, Halodule Wrightii (Shoal Grass) University of Texas Rio Grande Valley ScholarWorks @ UTRGV Biology Faculty Publications and Presentations College of Sciences 5-2016 Flowering and seed production in the subtropical seagrass, Halodule wrightii (shoal grass) Joseph L. Kowalski The University of Texas Rio Grande Valley Hudson R. DeYoe The University of Texas Rio Grande Valley Follow this and additional works at: https://scholarworks.utrgv.edu/bio_fac Part of the Biology Commons Recommended Citation Kowalski, Joseph L. and DeYoe, Hudson R.. "Flowering and seed production in the subtropical seagrass, Halodule wrightii (shoal grass)" Botanica Marina, vol. 59, no. 2-3, 2016, pp. 193-199. https://doi.org/ 10.1515/bot-2015-0099 This Article is brought to you for free and open access by the College of Sciences at ScholarWorks @ UTRGV. It has been accepted for inclusion in Biology Faculty Publications and Presentations by an authorized administrator of ScholarWorks @ UTRGV. For more information, please contact [email protected], [email protected]. DE GRUYTER Botanica Marina 2016; 59(2-3): 193–199 Short communication Joseph L. Kowalski* and Hudson R. DeYoe Flowering and seed production in the subtropical seagrass, Halodule wrightii (shoal grass) DOI 10.1515/bot-2015-0099 and Kowalski 2014). Halodule wrightii Ascherson (shoal Received 26 November, 2015; accepted 14 April, 2016; online first 25 grass) did not experience overall decline and remains an May, 2016 abundant and ecologically important seagrass that often grows in disturbed sediments (Pulich 1982). Reproduc- Abstract: Reproductive structures of the seagrass, tive structures of H. wrightii have been found from Brazil, Halodule wrightii Ascherson (shoal grass) are cryptic several localities along the Gulf of Mexico and Caribbean and ephemeral, but were found on several occasions at Sea and the east coast of North America (Table 1). Like several sites over more than a decade in the subtropical most seagrass species, the flowers of H. wrightii are dioe- Laguna Madre (USA). Structures align with descriptions cious (Moffler and Durako 1987). The broad geographical for H. wrightii. Male, but no female flowers were found, range in which H. wrightii flowers and sets fruit suggests but the numerous fruit-bearing shoots imply the occur- that sexual reproduction of H. wrightii is more common rence of pistillate flowers. Mean seed bank density was and cosmopolitan than the literature indicates, but the 890 seeds m-2 (1900–5200 m-2). Sexual reproduction in cryptic and ephemeral nature of these structures hinders H. wrightii is more common than previously believed and documentation. The purpose of this investigation was to the seed bank reserve may contribute to its ecological role describe the occurrence, abundance, and selected areal as a colonizer of disturbed sediments. extent of sexually reproductive material of H. wrightii in Keywords: flowers; Halodule wrightii; reproductive the LM, to compare and update these observations with ecology; seed bank. literature accounts, and to address its taxonomic standing. We found flowering shoots, fruits, and seeds of the seagrass H. wrightii in LM during late spring and summer of 1995, 1996, 2007, 2008, and 2009 at three locations. Seagrasses grow and expand by clonal expansion, estab- Prior to these observations, Thalassia testudinum Banks lishment of vegetative fragments, and sexual reproduction ex König (turtle grass) and Syringodium filiforme Kützing (Hall et al. 2006). Of these mechanisms, sexual reproduc- (manatee grass) were the only seagrass species docu- tion is critical to maintain the genetic diversity of popu- mented to have produced flowers and fruit in the LM lations (Reusch et al. 2005). The presence of flowers and (Kaldy and Dunton 1999). We define “fruits” as pollinated seeds is also important in the taxonomic description of pistillate flowers attached to the parent shoot irrespective a species (Phillips 1960). Seeds of some ruderal seagrass of stage of development, and “seeds” as detached from species can also play a key role in revegetation of dis- the parent shoot. Sediment seed reserve is defined here as turbed seagrass beds (Rasheed 2004). Three quarters potential reproductive units, irrespective of viability. (ca. 500 km2) of the sea bed of the subtropical lagoon, Discovery of H. wrightii seeds was initially made ser- Laguna Madre (LM) of Texas has historically been vegetated endipitously as part of a study of leaf production tech- (Onuf 2007), but an extensive freshet in 2010 resulted in niques for H. wrightii (Kowalski et al. 2001) performed decreased areal cover in most seagrass species (DeYoe between 1995 and 1997. Reproductive shoots and seeds were found in cores (9-cm diameter, 10-cm deep) taken *Corresponding author: Joseph L. Kowalski, Center for Subtropical for biomass and shoot production determinations (Kow- Studies and Department of Biology, The University of Texas Rio alski et al. 2001). Flowers, fruits, and seeds were found Grande Valley, 1201 West University Drive, Edinburg, TX 78539, USA, from June 1995 through September 1995 and from June e-mail: [email protected] 1996 through August 1996 at LM107 (Table 2) on nine sam- Hudson R. DeYoe: Center for Subtropical Studies and Department of Biology, The University of Texas Rio Grande Valley, 1201 West pling dates. Over the first four sampling dates in 1995, University Drive, Edinburg, TX 78539, USA 24 cores were collected. Eight cores were collected from 194 J.L. Kowalski and H.R. DeYoe: Flowering and seed production in production seed and Flowering H.R.DeYoe: and J.L. Kowalski Table 1: Record of previous occurrences of flowers, fruits, and seeds in Halodule wrightii from different localities. Locality Reproductive Comments Timing/Season Source structure(s) Texas, USA (Redfish Bay) Flowers (male and Fruit maturation in laboratory culture Apr McMillan (1976) female), fruit Texas, USA (Redfish Bay and Port Mansfield, Seeds Seeds sieved from sediment (seed bank estimates) No data McMillan (1981) Lower Laguna Madre) Download Date|7/7/168:01 PM St. Croix, US Virgin Islands (Jacks Bay) Seeds Seeds sieved from sediment (seed bank estimates) No data St. Croix, US Virgin Islands (Jacks Bay) Flowers (male and Early Apr maximum staminate production Mar-early May Johnson and female), fruit Maximum fruit production in mid-Apr Williams (1982) Unauthenticated Brazilian coast (several localities) Flowers (male and Taxonomic/descriptive No data Oliveira et al. female), fruit (1983) Texas, USA (Upper and Lower Laguna Madre) Seeds Seeds sieved from sediment (seed bank estimates) No data McMillan (1985) Alabama, Louisiana, Mississippi (Mississippi Flowers (male and 65% of flowers staminate Late May–June Eleuterius (1987) and Chandeleur Sounds) female?) North Carolina (Pamlico, Bogue, Roanoke, and Flowers (male and Salinity 12–34 May–Aug (flowers) Ferguson et al. wrightii Halodule Core Sounds) female), fruit, seeds (1993) Mississippi Sound (Alabama) Fruit – Late June–Sept McGovern and Blankenhorn (2007) Texas, USA (Lower Laguna Madre – LLM107) One male flower, See Kowalski et al. (2009) for site description June–Sept This study (26° 09′ N, 97° 14′ W) fruit, and seeds Site ABC: Approximately 5 km north of the Fruit 50–70 cm water depth June–July This study Brazos-Santiago Pass, west of Andy Bowie Park Thirteen male 50–70 cm water depth June–July This study (26° 08′ N, 97° 10 W) flowers and fruit Site IB (Isla Blanca Park, South Padre Island, Texas): Approximately 1 km north of the Brazos- Santiago Pass (26° 04′ N, 97° 09′ W) DE GRUYTER DE GRUYTER J.L. Kowalski and H.R. DeYoe: Flowering and seed production in Halodule wrightii 195 Table 2: Number of flower-bearing and fruit-bearing shoots, individual detached seeds, and percentage of sexually reproductive shoots sampled from 9-cm diameter (0.006 m2) cores for Halodule wrightii from LLM107 (1995–1996), and hand-scooped sediment samples from sites ABC and IB (see Table 1 for details and locations) from 2007 to 2009, Lower Laguna Madre, Texas. Sample date % cores w/ flowers Flower-bearing Fruit-bearing Detached seeds % Sexually Sediment seed or fruits shoots (per core) shoots (per core) (per core) reproductive shoots reserve (m-2) 24-June-95a 38 0 16 25 4.23 4064 7-July-95a 4 0 2 3 8.70 487 18-July-95a 42 0 18 32 4.99 5201 5-Aug-95a 28 0 3 3 3.75 487 19-Aug-95a 4 0 1 0 2.78 0 2-Sep-95b 0 0 0 7 0.00 1138 29-Sep-95b 0 0 0 2 0.00 325 4-May-96b 0 0 0 0 0.00 0 15-June-96b 38 0 8 9 6.11 1463 27-July-96b 62 0 9 12 4.05 1950 24-Aug-96b 12 1 1 0 3.30 0 23-June-07c NA 0 NA 6 NA NA 28-June-08c NA 0 NA 7 NA NA 13-Apr-09c NA 13 NA NA NA NA Percentage of sexually reproductive shoots based on the ratio of flower and fruit-bearing shoots to total shoot count per core. a24 cores collected per sampling date. bEight cores collected per sampling date. cNot based on cores (haphazard scoops of sediment). the site on all subsequent trips (Kowalski et al. 2001). and from 28° to 31°C during seed production (June to Sep- Reproductive structures on shoots were noted during pro- tember) while salinity varied from 32 to 40. The production cessing and enumerated. Length and width of staminate of flowers and fruits in tropical and subtropical seagrasses, flowers and length and diameter of fruits and seeds were including H. wrightii, has been correlated with water tem- measured to the nearest 0.1 mm using a caliper. Fruits were perature and, to a lesser degree, photoperiod and occurs typically found in pairs attached to maternal shoots. Fifteen during specific times of the year (McMillan 1976, 1982). A fruits were measured. Thereafter, individual H.
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